Bulletin of the Museum of Comparative Zoology at Harvard College

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HARVARD UNIVERSITY

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Library of the

Museum of

Comparative Zoology

•

BULLETIN

OF THE

MUSEUM OF COMPAEATIVE ZOOLOGY

AT

HARVARD COLLEGE, IN CAMBRIDGE

VOL. 117

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

The Cosmos Press, Inc.
Cambridge, Mass., U.S.A.

J

CONTENTS

PAGE

No. 1. — A Study op the Sharks of the Suborder Squa-
LOIDEA. By Henry B. Bigelow and William C.
Schroeder. (4 plates) Augrust, 1957 ... 1

No. 2. — Check List of the Reptiles and Amphibians
OF East Africa (Uganda ; Kenya ; Tanganyika;
Zanzibar). By Arthur Loveridge. August, 1957 151

No. 3. — The Spider Genera Crustulina and Steatoda
IN North America, Central America, and the
West Indies (Araneae, Theridiidae). By Herb-
ert W. Levi. August, 1957 . . . ' . .365

No. 4. — Contribution to a Revision of the Earthworm
Family Ocnerodrilidae. The Genus Nemato-
GENiA. By G. E. Gates. September, 1957 . . 425

No. 5. — Triassic Reptile Footprint Faunules from Mil-
ford, New Jersey. By Donald Baird. (4 plates)
November, 1957 447

BuUetin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 117, No. 1

A STUDY OF THE
SHARKS OF THE SUBORDER SQUALOIDEA

By Henry B. Biqelow and William C. Scheoeder

With Four Plates

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

August 1957

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Bulletin of the Musetun of Comparative Zoology

AT HARVARD COLLEGE
Vol. 117, No. 1

A STUDY OF THE
SHARKS OF THE SUBORDER SQUALOIDEA

By Henry B. Bigelow and William C. Schroeder

With Four Plates

CAMBRIDGE, MASS., U. S. A.

PRINTED FOR THE MUSEUM

August 1957

No. 1 — A Study of the Sharks of the Suborder Squaloidea '
By Henry B. Bigelow and William C. Sohroeder

TABLE OF CONTENTS

PAGE

Foreword 5

Acknowledgments 5

Classification of the Squaloidea 7

Key to Families of Squaloidea 13

Family Oxynotidae 13

Genus Oxynotus 14

Key to species of Oxynotus 17

Family Squalidae 17

Key to Subfamilies of Squalidae 18

Subfamily Squalinae 18

Synopsis of Genera of Squalinae 24

Genus Squulu^ 26

Genus CirrMgaleus 37

Genus Centroscyllium 38

Provisional Key to Species of Centroscyllium 47

Genus Etmopterus 47

E. h ullisi n. sp 50

Key to Species of Etmopterus 60

Genus Centropliorus 63

Nomenclatural status of the type species 73

C. foliaceus Giinther 1877 78

Key to species of Centropliorus 83

1 Contribution 848, Woods Hole Oceanographic Institution.

4 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus Centroscymnus 84

Key to species of Centroscymnus 95

Genus Scymnodon 96

Key to species of Scymnodon 101

Genus Deania 101

Key to species Deania 107

Subfamily Dalatiinae 109

Key to Genera of Dalatiinae 112

Genus Dalatias 113

Genus Somniosus 115

Key to species of Somniosus in the Northern Hemisphere 122

Genus Isistius 123

Genus Scymnodalatias 124

Genus Euprotomicrus 126

Genus Squuliolus 128

Genus Keteroscymnoides 132

Subfamily Echinorhininae 134

Genus Echinorhinus 134

References 136

BIQELOW AND SCHROEDER : SQUALOID SHARKS 5

FOREWORD

The sharks that lack an anal fin but do have snouts of the
ordinary "shark" form, and pectoral fins that are wholly pos-
terior to the gill openings (grouped together here as the sub-
order Squaloidea), have been surveyed of late for Philippine
waters and adjacent seas by Fowler (1941) and for the western
North Atlantic by Bigelow and Schroeder (1948). But the most
recent comprehensive accounts of the group for the oceans as a
whole, by Regan (1908) and by Garman (1913, pp. 188-244),
now lie many years in the past. And so much additional in-
formation has come to hand during these years, from various
sources, that the time seems ripe for a fresh survey of the group.
Many taxonomic questions, however, still await for their solution
the critical examination of large series of specimens. We attempt
little more in the following pages than to summarize the present
state of knowledge of the group. Anything as ambitious as would
be implied by the word "revision" is a task for the future.

It will save repetition if we state here (once and for all) that
the names of the species we have seen are preceded by an asterisk

Also, we should remind the reader, who may not be familiar
with the use of the trawl in deep water, that the depths of cap-
ture reported by this method, under the several genera, are in
reality those at which the trawl was dragging on the bottom.
This is not necessarily the depth at which the shark in question
was living, for it is always possible that any particular specimen
(even of the species that hold closest to the bottom) may have
been picked up when the trawl was being lowered, or hauled up
again through the water. The only precise data in this regard
are for specimens caught on hook and line. And we have very
few definite records of depths of capture for line-caught sharks.

ACKNOWLEDGMENTS

We welcome the opportunity of thanking our colleagues in
different parts of the world who have assisted us in various ways :
especially Dr. Tokiharu Abe, Tokyo, for contributing a specimen
of Somniosus long us; the Australian Museum, through Dr. G.
P. Whitley, for a specimen of Oxynotus hruniensis; Dr. G. Bel-
loc of the Museum, Monaco, for the gift of an excellent specimen

6 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

of Oxynotus from Italy; Dr. James Bohlke, of the Academy of
Natural Sciences for making available to us for study the type
specimen of Heteroscymnoides marleyi, from the Natal Coast,
southern Africa; the British Museum for the loan of the speci-
mens of C entroscymnus crepidater and of Scymnodon ringens,
on which Giinther (1870) based his descriptions of these species,
and of a specimen of Oxynotus paradoxus ; Dr. Paul Budker,
of the Paris Museum, for denticles from the type specimen of
Scymnodon ohscurus, and for drawings of its teeth; Dr. Fer-
nando Frade for re-examining for us the type specimen of Oxy-
notus paradoxus; Dr. Arni Fridriksson for specimens of
Centrophorus sqiiamosus and of Somniosus microcephalus from
Iceland; Dr. J. A. F. Garrick for kindly allowing us to quote
from his manuscripts in advance of publication, for a specimen
of a New Zealand shark apparently not separable from Centro-
scymnus crepidater of the eastern North Atlantic and for a para-
type of Etmopterus abernathyi; Dr. Finnur Gudmundsson, of
the Museum of Natural History, Reykjavik, for the loan of an
excellent series of C entroscymnus crepidater (see p. 91) ; Col.
John K. Howard for specimens of Squalus megalops from New
South Wales; Mr. N. B. Marshall for notes on Euprotomicrus
(p. 127) ; Dr. George S. Myers, of the Natural History Museum,
Stanford University, for loaning us a paratype of Centroscyllmm
ruscosum; Mr. G. Palmer, of the British Museum, for contribut-
ing tracings of pectoral fins of Richardson's young specimen of
Acanthias vulgaris, renamed Squalus megalops by Regan; Dr.
J. R. Pfaff, University Zoological Museum, Copenhagen, for the
loan of specimens of Centrophorus squamosus, Deania calcea, and
Centroscymnus crepidater, from Icelandic waters, and for the
gift of a specimen of C. crepidater; Dr. A. M. Ramalho for many
kindnesses, including the contribution of a series of Oxynotus
centrina from off Portugal and of an excellent specimen of Echi-
norhinus hrucus from northwest Africa, also for photographs of a
specimen (now in the Bocage Museum in Lisbon) that had been
described and pictured in 1870 by Capello under the name Lae-
margus rostratus Risso 1826; Dr. Leonard Schultz, for making
available to us the extensive collections of the U. S. National
Museum, as well as for assistance in various other ways; Dr. J.
L. B. Smith, Rhodes University, Grahamstown, for photographs
of the second dorsal fin spine of the shark described by Gilchrist

BIGELOW AND SCHROEDER : SQUALOID SHARKS 7

1922 as Atractophorus armatus; Dr. Ethelwyn Trewavas of the
British Museum for contributing a drawing of a lower tooth of
the type specimen of Centroscymnus macracanthiis as well as for
constant cooperation; Mr. Denys W. Tucker of the British Mu-
seum for drawings of the first and second dorsal fins of the type
specimen of Centroscymnus macr acanthus ; and Dr. Gilbert P.
Whitley, of the Australian Museum, for making available to us a
lower tooth from the type specimen of Centrophorus harrisonii
and for contributing a tracing of the pectoral fins of the type
specimen of Squalus megalops.

Drawings are by Eugene N. Fischer, Jessie H. Sawyer and
Henry B. Bigelow, except as otherwise noted. Photographs were
contributed by the Woods Hole Oceanographie Institution.

CLASSIFICATION OF THE SQUALOIDEA

While the sharks in question are an extremely homogeneous
assemblage differing conspicuously in their general makeup from
the other groups of sharks with which they share the lack of an
anal fin (pristiophoroids and squatinoids), the taxonomic rank-
ing assigned to them by recent authors has reflected divergent
viewpoints. Thus they have been grouped with the pristiophor-
oids by Garman (1913, pp. 12, 13) as the "group of families"
Squaloidei; by Berg (1940; Eng. Trans. 1947, p. 381) as the
suborder Squaloidei; by White (1937, pp. 100, 101) as the sub-
order Squalida; and by Bertin (1939, pp. 17-19) as the suborder
Squaliformes. Fowler (1941, pp. 222, 279) has placed the
squaloids in a separate order (Cyclospondyli), joining the
pristiophoroids with the squatinoids as the order Squatinae.
Bigelow and Schroeder (1948, p. 77) classed the squaloids as
one suborder (Squaloidea), the pristiophoroids as a second sub-
order (Pristiophoroidea), and the squatinoids as a third sub-
order (Squatinoidea) of the order Selachii. Still more recently
Berg, 1955, while similarly classifying the squaloids and the
squatinoids as suborders, has set the pristiophoroids apart as a
separate order. And this arrangement has much to recommend
it for the pristiophoroids are distinguished not only by their saw-
like snout, but l3y the "presence of a separate antorbital bar,
from which the upper jaw is suspended by a broad ligament, in
addition to the articulation to the cranium" (Bigelow and
Schroeder, 1948, p. 532).

8 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Characters. Sharks without an anal fin, but with the snout of
the ordinary shape (i.e. not beak-like and without lateral teeth
or cirri), and subcylindrical in general form, the anterior mar-
gins of the pectoral fins not expanded forward past the first pair
of gill openings. Two dorsal fins, with or without fin-spines.
Only 5 gill openings, all of them anterior to the pectorals. Inner
margins of pelvics entirely separate posterior to cloaca. Nostrils
entirely separate from mouth. Spiracles present and conspic-
uous. Eyes without nictitating fold or membrane. (For internal
characters, see Bigelow and Schroeder, 1948, p. 449.)

Families. Views have differed as to whether a subdivision of
the Squaloidea into separate families is justified on any basis
that has yet been proposed. Garman (1913, p. 13) distributed
them among three families: Squalidae for those with spines in
the dorsal fins, Scymnorhinidae and Echinorhinidae for those
without spines, separating these last two, one from the other, by
differences in their teeth. And this same scheme was accepted
by Bigelow and Schroeder (1948, p. 450), except that they re-
placed the name Scymnorhinidae by Dalatiidae because the gen-
eric name ScymnorJiinus Bonaparte 1846 is synonymous with
Dalatias Rafinesque 1810. Bertin (1939), however, recognized
only two families among the sharks in question — Squalidae for
those with fin spines, and Scymnorhinidae for those without
spines — while Berg (1940) has united them all in the single
family Squalidae. And Hubbs and McHugh (1951, pp. 163, 164),
for the reasons stated on p. 9, follow Berg in their suggestion
"that the Dalatiidae be fused with Squalidae" and that the
Echinorhinidae "should likewise be synonymized with Squali-
dae," though they retain the Dalatiinae and Echinorhininae as
subfamilies, pending future study.

These differences do not stem from any recent increase in
factual knowledge, for the external features of the squaloid
sharks were about as well known structurally a hundred years
ago as they are today. What the different systems do chiefly
mirror is the opinions held by the several authors as to the de-
grees of consanguinity among the animals in question. The ex-
pression, however, of views in this regard is but the secondary
purpose of animal classification; the primary purpose, as so
trenchantly worded by Simpson (1945, p. 13), is to "provide
a convenient practical means by which zoologists may know what

BIQELOW AND SCHROEDER : SQUALOID SHARKS 9

they are talking about and others may find out." In the ease
of any suborder as homogeneous structurally as the squaloid
sharks are, and for which the fossil record throws little light on
their evolutionary history, the decision, whether any subdivision
into different families is warranted depends primarily (in our
opinion) on whether it will help anyone from the practical
standpoint. The scheme adopted here is based on this premise.
At the same time it also emphasizes the strong probability that
one of the genera with which we are here concerned (Oxynotus)
represents a separate line of descent, for it is set apart from all
the others not only by the arrangement of its upper teeth, by the
nature of its dermal denticles, and by its body form (p. 14), but
also because it was differentiated at least as early as the Miocene.

Oxynotus was made the basis of a separate family (Oxynoti-
dae) by Jordan (1923, p. 103), followed by Key (1928, p. 467).'
And this course is followed here, as better indicative of the ap-
parent degrees of genetic relationship among squaloid genera
than was our former inclusion of it in the family Squalidae
(Bigelow and Schroeder, 1948, p. 450).

The Linnean genus Squalus, which includes the common spiny
dogfishes, has been regarded as what may be termed the "key
genus" among the squaloids that remain after the subtraction
of Oxynotus. This implies that the ancestors of the modern
squaloids had spines in their dorsal fins. And this view, recently
urged by Hubbs and McHugh (1951), is at least consistent with
the fossil record, for Protospinax Woodward 1919, from the
Upper Jurassic of Bavaria, with a spine of the squaloid type
in each of its dorsal fins, but with a small anal fin (if Wood-
ward's interpretation of the specimen be correct), and with
teeth but little differentiated in nature, may in some measure
bridge the gap between the squaloid sharks and the galeoids.
Hubbs and McHugh further argue that the absence of a spine
in the second dorsal fin in such of the sharks in question as lack
one (also of a spine in the first dorsal fin in all but one of the
genera in question) probably does not point to any close relation-
ship among them, but more likely represents the end result
of convergent evolution (i.e. loss of fin spines) along sep-
arate phyletic lines leading back to as many separate spiny-
finned ancestors. As one illustration, they point out that "com-

2 Regarded as a subfamily by Fowler, 1941, p. 223.

10 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

plete obsolescence of the spines would render Centroscymnus
scarcely separable from Somniosiis." This leads them to con-
clude that the presence or absence of spines in the dorsal fins has
been given undue weight in current classifications of the squaloid
sharks (as by Bigelow and Schroeder, 1948) in which it is in-
voked as the basis for the separation of families. Here the fossil
record does not help us at all, for this reaches as far back for the
genera Dalatias and Isistius, which lack fin spines, as for the
spiny-finned genera Squalus, Etmopterus and Centrophorus, i.e.
to the Upper Cretaceous — always provided that identification of
the fossils in question with these modern genera was correct.
For that matter, a shark with a rostral blade (but lacking rostral
teeth) that may reasonably be regarded as ancestral to the
pristiophoroids (Woodward 1932, p. 476, PI. 18; see also Jaekel,
1890, for other early pristiophorid fossils) is similarly known
from the Upper Cretaceous ; Sqnatina is even known from as far
back as the Jurassic. Thus both the squaloids with spiny fins
and those lacking spines, the progenitors of the pristiophoroids
with their bizarre snouts, and the squatinoids (ray-like in ap-
pearance though true sharks) had completed their major evolu-
tionary history by the Upper Cretaceous period, which (the
geologists tell us) lies something like 75 million years in the past.
For all that is known to the contrary, this may equally apply to
the aberrant squaloid genera Oxynotus and Echinorhinus, though
the known record for these runs back only to the Miocene. (For
the fossil record of modern shark genera, see Romer 1945, pp.
576-577.)

Were one approaching the problem anew, the shape of the
caudal fin, combined with the degree of elevation of the caudal
axis, would seem about as appropriate as would the presence or
absence of fin spines for a primary alternative among the genera
of squaloids. Thus the one character is about as susceptible of
precise determination as the other, while, in each case, the ex-
treme states are bridged by intermediates so that division is not
sharp-cut on either basis. Also the use of either one would result
in the same grouping of genera as would the use of the other,
with one exception.

A third character that is alternative between the groups of
genera under discussion here, though it seems not to have been
emphasized previously in this particular connection, is the con-

BIGELOW AND SCHROEDER : SQUALOID SHARKS

11

i^sSSfeii -(iSsJi^^S^

Fig. 1. Corner of mouth, of different squaloids, to show preoral clefts and
preoial pouches, if any; the latter outlined by the broken curves. A,
Oxynotus centrina, male 590 mm. long, Mus. Comp. Zool. No. 3!;»576, x about
3. B, Centrophorus squamosiis, female 1230 mm. long, Iceland, Mus. Comp.
Zool. No. 37825, x about 0.5. C, Centroscymnus coelolepis, female, 1035
mm. long, offing of Delaware Bay, Mus. Comp. Zool. No. 38295, x about 2.
D, Centroscymnus crepidatcr, female, 784 mm. long, Faroes, Mus. Comp.
Zool., No. 39377, with preoral cleft and upper labial furrow pinned open, x
about 1. E, Dalatias Helm, male, 1114 mm. long, Japan, Mus. Comp. Zool. No.
1116, X about 0.5. F, Isistius hrasiliensis, female, 501 mm. long, Japan, Mus.
Comp. Zool. No. 1245, x about 2.

12 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

formation of the more or less voluminous inpocketing of the
lower surface of the head, that bounds each angle of the jaw
on the outer side. In all the known genera of Squaloidea that
have a spine in the second dorsal fin as well as in the first (in-
cluding Oxynotus, Fig. lA), this oral pocket extends inward-
forward across the lower surface of the head, anterior to the
upper jaw, as an open and conspicuous preoral cleft, that is
simple in some (e.g. Oxynotus, Fig. lA; Centroscymnus crepi-
dater, Fig. ID), but that expands inwardly in most to form a
more or less voluminous subdermal pouch (e.g. in Centroscymnus
coelolepis, Fig. IC). On the other hand, the majority of the gen-
era that lack a spine in the second dorsal fin equally lack the open
preoral cleft, although the preoral pouch extending forward from
each of the oral pockets is voluminous (Fig. IE, F). Two, how-
ever, of the genera of this group, Scymnodalatias and Echino-
rhinus do have short preoral clefts.

And, in any ease, the group difference in this respect, while a
convenient aid in the definition and identification of genera, is
obviously of the kind rated by White (1937, pp. 51-53) as of
''physiological importance," rather than as an indication of
lines of ancestral descent.

There seems, in short, to be "no securely objective basis," as
pointed out by Hubbs and McHugh (1951, p. 163) "on which
to choose between the characters to be emphasized" in the classifi-
cation of the Squaloidea.

But the cause of stability in nomenclature seems best served
by using the presence or absence of fin spines as the primary
alternative, since this accords with precedent, whereas to use
the shape of the caudal fin, as have Hubbs and McHugh (1951,
p. 164) in their analytical key to the dalatiine sharks, or to
emphasize the presence or absence of the preoral clefts, would re-
sult in a novel alignment which we could not support by any
cogent reasoning. But we wish it understood that this choice
does not carry any phylogenetic implication but is purely a mat-
ter of convenience.

In our earlier synopsis of the Squaloidea (Bigelow and
Schroeder 1948, p. 450), we recognize two families for such of
the genera as lack a fin spine in the second dorsal fin : Dalatiidae
for those in which the teeth have only one cusp, but are of con-
spicuously different shapes in the two jaws, contrasted with

BIGELOW AND SCHROEDER : SQUALOID SHARKS 33

Ecliinorhinidae (for Ecliinorhinus), in which the teeth are simi-
lar in the two jaws, with several cusps. But it seems more logical.
in view of the uncertainties just discussed, to follow Hubbs and
McHugh (1951) in reducing these categories to the rank of sub-
families of the family Squalidae, the structural contrast between
them obviously being much narrower than that between either
of them and the genus Oxynotus (p. 14).

Key to Families of Squaloidea

Only two or three upper teeth in the hrst row, but with the number of
teeth greater by one in each successive row, so that the rows of func-
tional teeth are successively longer from front to rear; trunk sub-
triangular in cross section with a prominent ridge low down on either

side anterior to the pelvic fins Family Oxynotidae, p. 13

About as many upper teeth in the anterior row as in the succeeding
rows, so that any one functional row is about as long as any other;
trunk subcylindrieal in cross section in most; longitudinal ridges, if

any, confined to the tail sector posterior to the pelvic fins

Family Squalidae, p. 17

Family OXYNOTIDAE

Family cliaracters (based on only known genus, Oxynotus).
Squaloidea with the upper teeth occupying a triangular area on
the roof of the mouth, in quincuucial arrangement, the functional
rows successively longer from front to rear, the first row con-
sisting of three teeth only (on specimen studied, two, according
to Eey 1928, p. 469), the number of teeth greater by one in each
successive row, and with about six rows functional; lower teeth
in a single functional row, blade-like, the median tooth sym-
metrically triangular, but the successive teeth increasingly
oblique, outward (PI. 1) ; the uppers smooth-edged, but the
lowers finely serrate. The trunk subtriangular in cross section,
with a prominent longitudinal ridge low down on either side an-
terior to the pelvic fins; preoral clefts well developed and very
conspicuous, extending forward-inward so far that they are sep-
arated by only a narrow isthmus in the midline of the snout ( Fig.
lA) ; lips thick, spongy, with a complex series of cross folds (Fig.

14 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

lA) ; first dorsal fin spine slanting either rearward or forward;'
the first dorsal fin either with or without supporting radial
cartilages posterior to the imbedded portion of the spine ; both fin
spines conspicuously stout, but projecting only slightly beyond
the skin. Dermal denticles so large and prominent that the skin
is very rough. On young specimens they are tridentate, with the
median point much the longest. But they are much more com-
plex in form on adults, mostly with three subsidiary points, the
basal point with a varying number of radial furrows (for excel-
lent illustrations of the denticles, see Rey 1928, p. 471, Fig. 157).

Genera. Only one genus is known : Oxynotus Rafinesque 1810.

Remarks. The pattern in which its upper teeth are arranged
sets Oxynotus apart from all other squaloid sharks — from all
other sharks for that matter — which of itself seems to us war-
rant enough for referring it to a separate family.

Genus OxYNOTUS Rafinesque 1810

Oxynotus Eafinesque, 1810, pp. 45, 60, type species *Squnlus oentrina Lin-
naeus 1758, p. 233, type locality, Mediterranean, For generic synonyms,
see Garman, 1913, p. 190.

Generic characters. Those of the family (p. 13). Maximum
length probably not far from one meter.

Depth range. The type species (centrina) has attracted scien-
tific attention at least since the time of Belon (1515, p, 46 bis),
and is well known to Mediterranean fishermen, in moderately
deep water. But the only definite statements we have found as
to depth of capture are Roule's (1919, p. 123) report of a
Mediterranean specimen taken at 60 meters, and Poll's (1951, p.
57) report of three, from 100-180 meters off equatorial West
Africa. Depth records for the newly discovered *0. paradoxus
(p. 16) have ranged between 265 and 530 meters (Tucker and
Palmer 1949, p, 930). All we have found for *hruniensis of the
Australian-New Zealand region is that a specimen at hand
(p. 5) was trawled at about 180 meters.

3 This seems an appropriate place to remind the reader that it is only the
first dorsal fin spine that is inclined forward in any Ijnown species of Oxynotus,
not both of the fin spines, as is erroneously stated in our earlier Icey to the

fenera of Squalidae (Bigelow and Schroeder 1948, p. 450, alternative lA) ; also
hat in constructing that l<ey we had overlooljed the conditions existing in
•0. paradoxus.

BIGELOW AND SCHROEDER : SQUALOID SHARKS

15

Fig. 2. A, dissection of first dorsal fin of Squalus acantliias, female,
about 800 mm. long, coast of Massachusetts, Mus. Comp. Zool. No. 458,
X about 0.5, to show radial cartilages. B, similar dissection of second dorsal
fin of same. C, similar dissection of first dorsal fin of female Oxynotus
centrina, 640 mm. long, Mediterranean (Mus. Comp. Zool. No. 39564), x
about 0.5; D, similar dissection of second dorsal fin of same, x about 0.5;
E, Oxynotus centrina, male, 590 mm. long, Mus. Comp. Zool. No. 39576,
upper jaw and t«eth, viewed from in front, x about 3.3; F, lower teeth of
same, functional row, viewed from outside the mouth, x about 3.8.

10 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Remarks. Previous to 1929 it had been generally agreed that
the forward inclination of the first dorsal fin spine in the two
species of Oxynotus that had been described up to that time
was (with the arrangement of the upper teeth) diagnostic of the
genus Oxynotus, which still remains the only known representa-
tive of the family Oxynotidae. But Frade's discovery in 1929
that the first dorsal spine in his new species *0. paradoxus, from
the coast of Morocco, is inclined rearward (see also Frade 1932
and Norman 1932) has shown that the direction in which it
slopes can no longer be regarded as a generic alternative here.

Species. Three species are known: (1) *centrina (Linnaeus)
1758, familiar to fishermen in the Mediterranean and neighbor-
ing parts of the eastern North Atlantic, ranging northward
regularly to the Bay of Biscay, also known as a stray to Cornwall
(Day 1880-1884, p. 319) and to Norway (Jenkins 1925, p. 321) ;
(2) *hruniensis (Ogilby) 1893, originally described from Tas-
mania and subsequently reported from southern Australian
waters and from New Zealand; and (3) * paradoxus Frade 1929,
eastern North Atlantic between Morocco and Ireland. For excel-
lent accounts and illustrations of the newly discovered *0. para-
doxus, see Frade 1932 and Norman 1932.

The most conspicuous respects in which these three differ ex-
ternally one from another are in the shapes of the dorsal fins
and in the direction in which the first dorsal spine is slanted.
For details we refer the reader to Norman's (1932) illustration
and key. A more interesting morphologic difference is that while
in * paradoxus and in *hru7iicnsis the first dorsal fin posterior to
the spine is supported, as in Squalus (Fig. 2A, B), by a series
of radial cartilages, in *centrina (Fig. 2C, D) the first dorsal
fin has no radial cartilages but is supported solely by the spine.

The dentition also differs in detail from species to species in
Oxynotus, as is so commonly the case in other genera of sharks.
Thus the upper teeth, which are narrow-awl-shaped in the cen-
tral rows but broadly triangular and blade-like in the outer rows
in *centrina (PI. 1), are relatively broader in the central rows
but relatively narrower in the outer rows in ^'paradoxus. Also,
the lower teeth differ in number from species to species, there
being 9 series in ^centrina (6 specimens counted) contrasted
with 13 in the type specimen of ^paradoxus, as Dr. Frade writes
us, and 13 (perhaps 15) in the one specimen of the latter that we

BIGELOW AND SCHROEDER : SQUALOID SHARKS 17

have examined (counting is difficult). Rey (1928, Fig. 156, p.
470), it is true, credits *centrina with 15, and pictures it thus.
But this was prior to the discovery of the species *paradoxus.
There are 13 series of lower teeth in the specimen of *hruniensis
we have at hand.

Key to Species of Oxynotus

1. First dorsal fin spine sloping a little rearward; distance from tip of
second dorsal spine to apex of second dorsal fin is about 1^^ times as
long as the spine, tip to base *paradoxus Frade 1929

Eastern North Atlantic, known off

the coasts of Morocco and of

Ireland, and from the Gulf of

Gascony. For records of occurrence,

see Krefft 1955 and Tucker and

Palmer 1949.

First dorsal fin spine sloping forward; distance from tip of second

dorsal spine to apex of second dorsal fin is not longer than the spine,

tip to base 2

2. Interspace between first and second dorsal fins is at least l^o times
as long as base of second dorsal; distance from tip of first dorsal fin
spine to apex of first dorsal fin is only V^-'/s as long as the spine, tip
to base; most, at least, of the dermal denticles on the sides with 3
cusps only *centrina (Linnaeus) 1758

Eastern North Atlantic, including

the Mediterranean.

Interspace between first and second dorsal fins little if any longer

than base of second fin; distance from tip of first dorsal spine to

apex of first dorsal fin is a little longer than the spine, tip to base;

most, at least, of the dermal denticles on sides with 5 cusps

*bruniensis (Ogilby) 189.3.
Australia, Tasmania, New Zea-
land.

Family SQUALIDAE

First dorsal fin, like the second dorsal, with a series of radial
supporting cartilages (Fig. 2, A, B) ; the trunk subcircular in
cross section in most (for an exception see under Cirrhigaleus,
p. 37) ; upper teeth as well as the lowers in several rows paral-
leling the jaw, with about as many teeth in any one row as in
any other; dermal denticles with one central cusp (if any) in

18 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

most, with several cusps in some (p. 18) ; preoral clefts present
in some, but lacking in others; see discussion, p. 12.

Key to Subfamilies of Squalidae

1. Each of the dorsal fins has a spine; preoral clefts present in all
known genera, see discussion, p. 12 ; lips smooth

Subfamily Squalinae,* p. 18
No spine in the second dorsal fin or in the first in most cases; preoral
clefts lacking in most genera but present in some (see discussion,
p. 12) 2

2. Teeth with only one cusp, the lowers much broader than the uppers ;
lips crenulate or smooth ; dermal denticles with only one cusp or spine

(if any) Subfamily Dalatiinae, p. 109

Teeth with several cusps, the uppers similar in shape to the lowers ;
dermal denticles often with two or more cusps or spines ; lips smooth . .

Subfamily Echinorhininae, p. 134

Subfamily SQUALINAE

Characters. Squalidae with a spine in each of the dorsal fins ;
the caudal axis raised, and the caudal fin wider below axis than
above ; longtitudinal dermal ridges, if any, confined to tail sector,
posterior to level of cloaca ; preoral clefts present in all known
genera.

Remarks. Fowler (1934, p. 239 ; 1941, p. 223) restricted the sub-
family Squalinae to those Squalidae in which the fin spines are
not laterally grooved, and in which the caudal fin does not have
a subterminal notch, to include Squalus Linnaeus 1758, and the
barbel-bearing Cirrhigaleus Tanaka 1912. For those with later-
ally grooved fin spines and caudal fin with subterminal notch, he
proposed Etmopterinae to include all the other known members
of the group. But this scheme results in the union (as Etmop-
terinae) of species in which the upper teeth have at least three
cusps (the lowers also in some) with other species in which the
uppers, like the lowers, have only one cusp, a dental difference

i Previous to 1896 the sharks in question had commonly been referred to as
Spinaeidae or as Spinaces ; e.g., by Miiller and Henle 1841, by Bocage and
Capello 186G. by Dtmieril 18G5, by Day 1880-1884, by Moreau 1881, and by
Smitt 1895. The "Centrophoroidei of Bleeker 1860 was an equivalent. The
substitution of Squalidae by Jordan and Evermann 1896 doubtless stemmed from
their realization that the generic name Spinax Cuvier 1817 was not available as
the basis tor the name of a family because it is a synonym of Etmopterus
Kaflnesque 1810.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 19

that would seem as sound a basis for subdivision as is the nature
of the fin-spines. And the use of the shape of the terminal part
of the caudal fin as a primary character here seems forbidden by
the differences in this respect among the species which (by their
dentition) fall within the genera Centroscymniis and Scymnodon.

Genera. The members of the subfamily Squalinae as here de-
fined fall in three divisions, as follows :

Division A — Fin spines without lateral grooves ; lower con-
tour of caudal fin without subterminal notch; teeth with only
one cusp, the cusps of the uppers as well as of the lowers so
strongly oblique as to form a nearly unbroken cutting edge
along the jaw ; the median tooth in each jaw basally overlapping
the tooth next to it on either side, and the base of each succeeding
tooth overlapping that of the next tooth outward along the jaw.
This division includes the genera Sqiialus Linnaeus 1758 and
Cirrhigaleus Tanaka 1912.

Division B — Fin spines with lateral grooves; lower contour
of caudal fin with subterminal notch ; upper teeth with three or
more cusps, not overlapping basally, the lowers similar to the
uppers in some (Centroscyllium) , but with only one cusp in
others {Eimopterus) , and overlapping basally as in division A.
This division includes two well-marked genera : Centroscyllium
Miiller and Henle 1841 and Etmopterus Rafinesque 1810.

Division C — Fin spines with lateral grooves and lower con-
tour of caudal with subterminal notch (at least in most cases)
as in division B; but teeth (uppers as well as lowers) with only
one cusp ; the uppers narrower than the lowers in most, either
loosely spaced or with their bases in contact or slightly over-
lapping, the lowers broad based, with each tooth overlapping the
next outward along the jaw as in division A. This group includes
all known members of the subfamily other than those listed above
for divisions A and B.

Students of sharks are now in general agreement as to the
alternative characters of the genera Sqiialiis, Cirrhigaleus (if
the latter be accepted as distinct from Squalus — see p. 37),
Etmopterus and Centroscyllium; i. e., divisions A and B. But
the picture is not yet clear for the members of division C, which
are deep-water species, few specimens of which have found their
way into museum collections.

20 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

The members of division C that were set apart the earliest from
the old genus Squalus Linnaeus 1758 were *'S. squamosus Bonna-
terre 1788, and *S. gramdosus Bloeh and Schneider 1801, both
of which were transferred by Blainville (1816, p. 121) to his new
genus Acanthorhinus, while squamosus was referred almost
simultaneously by Cuvier (1817, p. 130, footnote 3) to his Cen-
trina. But the earliest genus that was based in so many words on
a member of this division was Centrophorus, proposed by Miil-
ler and Henle in 1837, for *C. granulosus Midler and Henle,
which they considered identical with *8qualus granulosus Bloch
and Schneider 1801. And their subsequent description and illus-
tration of it (Mliller and Henle 1841, p. 89, PL 33) were so
satisfactory that there is no doubt as to the specific identity of
the Mediterranean specimen which they had at hand.

A year later Bonaparte (1838, p. 207; 1839, p. 9, extra) pro-
posed the generic name Lepidorhinus for *squamosus, while
Lowe in 1839 instituted Acanthidium for two Madeiran species,
one of which {*calceum Lowe 1839) falls in the group wdth which
we are dealing, whereas the other {pusillum Lowe 1839) is now
known to be an Etmopierus (p. 49).

Next, Gill (1862, pp. 496, 498) made the * Squalus mjato of
Rafinesque 1810 the basis of a new genus, Entoxychirus, setting
it off from the old genus Squalus by its fins, from Centrophorus
Miiller and Henle 1837 under the misconception that the teeth
of *iiyato are equally oblique in both jaws. This error was no
doubt based on an illustration that Bonaparte (1841, PI. 57,
fig. 2) credited to *uyato, but which actually pictured the teeth
of a Squalus, as Garman (1913, p. 198) has pointed out. The
relegation of Eyitoxychirus to the synonymy of Centrophorus by
Garman (1913, p. 196), but its revival by Fowler (1941, p. 242)
for species with "pectoral angle extended," contrasted with "not
or scarcely extended" in Centrophorus, covers the subsequent
history of the genus.

In 1864, Bocage and Capello proposed two new genera for
members of the division in question : Centroscymnus, type species
*C. coelolepis Bocage and Capello 1864, and Scymnodon, type
species *S. ringens Bocage and Capello 1864, based in each case
on a newly discovered shark from deep water off Portugal. These
species were described by them in greater detail, with excellent
illustrations two years later (Bocage and Capello 1866, pp. 30,

BIQELOW AND SCHROEDER : SQUALOID SHARKS 21

32, PI. 1, fig. 1, PI. 2, fig. 3, PI. 3, figs, 2a 2b). And while
Giinther (1870, p. 420) did not regard the dental differences on
which these two genera were based as sufficient to separate them
from Centrophorus, Garman (1906, p. 204) and Regan (1908, p.
39) revived them, Garman without stating his reasons, but Regan
because of the same dental characters on which Bocage and Ca-
pello had set them up originally.

Meantime, Johnson, in 1867, had proposed the genus Mache-
philus for his new Madeiran species dumerili, separating it from
Centrophorus because of the presence of a symmetrical median
tooth in its lower jaw. However, subsequent authors (Garman
1913, p. 212; Rey 1928, p. 440; Bigelow, Schroeder and Springer
1953, p. 224) have classed dumerili as a synonym of *Cen-
trophorus squamosus (Bonnaterre) 1788, i.e., have not con-
sidered the presence or absence of a symmetrical median lower
tooth as sufficient ground for generic separation, and this same
course is followed here (p. 72).

The next landmark in the taxonomic history of the squaline
sharks of division C was the institution by Jordan and Snyder in
1902 of the genus Deania, type species *eglantina Jordan and
Snyder, a Japanese shark characterized by a very long snout as
well as by pitchfork-shaped denticles on the sides of its trunk.
And this generic entity was accepted by Garman (1906, p. 205;
1913, p. 215). But Garman 's revival of the generic name Acan-
thidium Lowe 1839 for it, in place of Deania, was not well judged,
for the type species of Acanthidium {*pusillum Lowe 1839) as
designated by Goode and Bean (1895, p. 10), by Jordan and
Evermann (1896, p. 55), and by Jordan (1919, p. 195) falls
within the limits commonly accepted for the genus Etmopterus
Rafinesque 1810. Consequently, Fowler's (1941, p. 237) relega-
tion of Acanthidium to the synonymy of Etmopterus is to be
accepted — unless indeed Whitley's (1939, p. 266) separation of
Acanthidium, type species Centrina nigra Lowe 1834 {*Acan-
thiduim pusillum Lowe 1839^) from Etmopterus Rafinesque
1810, type species *Squalus spinax Linnaeus 1758, is to be fol-
lowed. And the grounds stated by Whitley (i.e. Acanthidium
with "first dorsal much smaller than the second and the ventrals
just before the level of the first dorsal") do not seem to justify

5 Whitley (1939, p. 266) points out that it was as Centrina nigra that Lowe
(1834, p. 144) first reported the shark which he renamed * Acanthidium pusillum
in 1839.

22 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

generic division in this case, for the relationship in size between
the first and second dorsals is about the same among our Nor-
wegian and Mediterranean specimens of *spinax (type species of
Etmopterus) as it is in the type species of Lowe's (1839) genus
Acanthidium. Also, the rear end of the bases of the pelvics ranges
in position from below the second dorsal spine to considerably
anterior to the latter among the several Atlantic species of
Etmopterus (Bigelow, Schroeder and Springer 1953, p. 272).

A year after the genus Deania had been proposed by Jordan
and Snyder (1902), Jordan and Fowler (1903, p. 633) made the
Japanese Centrophorus squamulosus Giinther 1877 the type of
still another genus, Zameus, separating it from Centrophorus by
the shape of its dermal denticles. And while Garman (1913, p.
207) classed Zameus unequivocally as a synonym of Scymnodon
Bocage and Capello 1864, Fowler (1941, p. 226) has revived it
as a subgenus of the latter, now invoking the length of the orbit
as compared with the length of the snout as the alternative sub-
generic character.

Next Smith and Radcliffe (1912, p. 681) made their new
Philippine species *profundorum the type of another new genus,
Nasisqualus, but without mentioning anything to separate it from
Deania Jordan and Snyder 1902, with which Regan (1912) has
synonomized it. And during the following year Garman (1913,
p. 206) proposed Centroselachus for the Portuguese shark that
Bocage and Capello (1864, 1866) had originally described and
pictured as *Centrophorus crepidater, basing its generic separa-
tion from Centrophorus Bocage and Capello 1864, and from
Scymnodon Bocage and Capello 1864 on small differences in the
shapes of the dermal denticles. Garman (1913, pp. 189, 211)
also revived the genus Lepidorhinus Bonaparte 1838, type species
"^Squaliis squamosals Bonnaterre 1788, but this revival has not
been accepted generally.

Nine years after the appearance of Carman's monograph, Gil-
christ (1922, p. 48, PI. 7, fig. 3) proposed the genus Atracto-
phorus for a southern African squalid, A. armatus Gilchrist, in
which the tip of the second dorsal fin spine is described as barbed,
like an arrowhead (p. 82). Next Whitley (1932, p. 326) pro-
posed Deaniops, Avith the Australian Acanthidium quadrispi-
nosum McCulloch 1915 as type, without including any generic
diagnosis at the time, but adding subsequently (Whitley 1939,

BIGELOW AND SCHROEDER : SQUALOID SHARKS 23

p. 266) that it Avas for "species with a long low first dorsal fin."
Two years later Fowler (1934, p. 293) proposed Proscymnodon,
type species Centrophorus plimketi Waite 1909, from New Zea-
land, as a subgenus of Scymnodon, because of length of orbit
combined with the positions of the first and second dorsal fins,
and Proscym7iodon has been accepted by Whitley (1934, p. 199;
1940, p. 142) as a full genus. Whitley (1940, pp. 145, 146)
subsequently has proposed two subgenera within the old genus
Centrophorus Miiller and Heule 1837, Gaboa (type species Cen-
trophorus harrisonii McCulloch 1915, Australia) and Somni-
spinax (type species Centrophorus nilsoni Thompson 1930, also
from Australia), the former because of "the form of its teeth
and denticles," the latter because of smooth edged teeth, "dif-
ferently shaped denticles, differently shaped snout, fins, etc.,"
but without more precisely specifying what these differences are.
As a final complication, Fowler's (1941, p. 223) definition of
Enfoxychirus as with "inner pectoral angles extended" corres-
ponds to Garman's (1913, p. 189) for Centrophorus, while
Fowler's for Centrophorus ("pectoral angles not or scarcely
extended") corresponds to Garman's for Lepidorhinus.

To sum up, such of the Squalinae as have grooved fin spines,
upper teeth as well as the lowers with only one cusp, and caudal
fin with a subterminal notch, have been referred to 18 different
genera or subgenera at one time or another. But our own study
of a broad representation of the group leads us to conclude that
the genera Centrophorus Miiller and Henle 1837, Centroscymnus
Bocage and Capello 1864, Scymnodon Bocage and Capello 1864,
and Deania Jordan and Snyder 1902, perhaps with the addition
of Atractophorus Gilchrist 1922, afford appropriate resting
places for all of them — provided that a genus, to be of practical
value, must be * ' separated from other similar units by a decided
gap" (Mayr, Linsley and Usinger, 1953, p. 48). And we may
add that we subscribe heartily to this proviso.

The characters that have proved the most useful, here, for the
definition of genera (because they are the most definitely alterna-
tive between groups of species) are: the shapes of the teeth, the
shape of the inner corner of the pectoral fins, the relative length
of the snout, the length of the exposed portion of the second dor-
sal fin spine, and perhaps also whether the tip of that spine is
smooth or is barbed or otherwise expanded, as it is described

24 BULLETIN : ilUSEUM OF COMPARATIVE ZOOLOGY

for one member of the group (p. 82). The differences in rela-
tive size, and in length at base, of the first and second dorsal fins
(sometimes invoked as of generic significance) are not sharply
alternative enough to be of much help in this regard. And recent
discoveries (p. 87) have also proved that the shape of the dermal
denticles on the sides of the trunk is not as reliable a generic
criterion here as we had formerly believed. For further details
in this last regard, see our earlier discussion (Bigelow and
Schroeder 1954, p. 47), to which we should add that as early as
1861 Steenstrup had already pointed out that the dermal den-
ticles of sharks do not persist throughout the life of the indi-
vidual, as the scales of bony fishes do, but are short lived, to be
replaced as they are shed by more newly formed denticles.

Synopsis of Genera of Squalinae

1. Upper teeth as well as the lowers with one cusp only 2

Upper teeth with 3-7 cusps 7

2. Fin spines well developed, Avithout lateral grooves, caudal fin without
subterniinal notch, its tip not truncate; upper teeth similar in shape

to lowers 3

Fin spines with lateral grooves ; lower margin of caudal fin with
a definitely outlined subterniinal notch and with its tip more or
less truncate, at least in most species ; upper teeth differing more or
less widely in shape from the lowers, except as noted on pages 26
and 38 4

3. Anterior margin of nostril does not bear a barbel ; caudal peduncle
with a longitudinal ridge along either side, and with a preeaudal pit
above but none below; upper teeth, like the lowers, with the successive
cusps directed outward strongly along either side of the jaw

Squalus Linnaeus 1758, p. 26
Anterior margin of nostril with a conspicuous barbel reaching beyond
the mouth; caudal peduncle without longitudinal ridges and without
preeaudal pit ; other characters, in general, including teeth, as in

Squalu.s Cirrhigaleus Tanaka 1912, p. 37

•4. inner corner of pectoral fins angular and more or less extended (see
discussion, p. 66) ; fin spines well developed, the second longer than
the first; upper teeth narrower than the lowers and more nearly erect
in most cases; lowers directed strongly outward; cutting edge of
lower teeth smooth in some but more or less irregularly and finely
serrate in others (p. 67, Fig. 9) ; snout in front of mouth not longer
than from mouth to level of origin of pectoral fins; dermal denticles on

BIGELOW AND SCHROEDER : SQUALOID SHARKS 25

sides of body low, block-like, scale-like, or conical

Centrophorus Miiller and Henle 1837, p. 63
Inner corner of pectoral fins not extended ; rounded in most, perhaps

quadrate in one kno-\vn species 5

Upper teeth midway along either side of the jaw noticeably longer
than those toward center of the mouth (Fig. 13C) ; first and second
dorsal fin spines projecting only slightly beyond the skin; dermal
denticles on sides of body low, scale-like, with tridentate rear margin;
other characters in general as in Centroscymnus (alternative 6; see

also p. 85) Scymnodon Bocage and Capello 1864. p. 96

Upper teeth midway along either side of jaw only slightly longer than

those toward center of mouth (Fig. 12C) 6

Snout in front of mouth not longer than from mouth to level of origin
of pectoral fins ; inner corner of pectoral fins broadly rounded ; dorsal
fin spines either exposed at the tip or entirely enclosed in the skin;
lower teeth smooth-edged, the cusps directed strongly outward; upper
teeth lancet-shaped, much narrower than lowers, and more nearly erect;
dermal denticles on sides of body scale-like, tridentate or evenly
rounded, low in adults of known species but rising more steeply from
skin on juveniles of at least one (p. 90) ; subterminal notch of caudal
fin well marked in most species, but indistinct in juveniles of one
named species, the parentage of which is not yet definitely known (p.

94) Centrosoymnus Bocage and Capello, 1864. p. 84

Snout in front of mouth longer than from mouth to level of origin of
pectoral fins; inner corner of pectoral fins rounded in most, perhaps
quadrate in one of known species (p. 102), but not extended at all;
both of the dorsal fin spines well exposed; caudal fin with well marked
subterminal notch; teeth smooth edged; the uppers narrow, triangular,
on broad bases; the lowers strongly oblique in the females of all
known species, but less so in the males of some (p. 103); dermal
denticles on sides of body pitchfork shaped, rising steeply from the

skin; other characters in general as in Centrophorus (p. 63)

Deania Jordan and Snyder 1902, p. 101
Lower teeth with 3-5 cusps and similar to the uppers in shape; dorsal
fin spines well developed, with lateral grooves; caudal fin truncate
and with subterminal notch in known species (but see discussion, p.
46) ; caudal peduncle without precaudal pits or lateral longitudinal

ridges; dermal denticles on sides of body slender, thorn-like

Centroscyllium Miiller and Henle 1837, p. 38
Lower teeth with only one cusp, much wider than uppers; dermal
denticles on sides of body bristle- or conical thorn-like to truncate;

other characters in general as in Cettti-oscyUivm

Etmopterus Rafinesque 1810. p. 47

26 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus SqUALUS Linnaeus 1758

Squalus Linnaeus 1758, p. 233, type species *S. acanthias Linnaeus, desig-
nated by Gill, 1861, p. 369.
For generic synonyms, see Bigelow and Schroeder, p. 452.

Generic characters. Squalinae with fin spines well developed,
without lateral grooves; caudal fin without subterminal notch
but with well marked lower-anterior lobe; caudal peduncle with
a longitudinal ridge along either side, and with a pre-caudal
pit above but none below ; margin of nostril without barbel ; pre-
oral clefts present, slightly expanded inwardly ; snout in front
of mouth considerably shorter than from front of mouth to origin
of pectorals; inner corner of pectoral fins either rounded, sub-
angular, or slightly extended ; dermal denticles lanceolate, heart-
shaped, or tridentate with sharp tip, and of different shapes on
different parts of the trunk as well as from species to species;
teeth in both jaws with only one cusp, directed so strongly out-
ward as to form a practically unbroken cutting edge along either
half of the jaw ; the base of each tooth overlapping that of the
next tooth, outward. Maximum length a little more than one
meter.

Depth range. The depth range recorded for this genus is
from the surface down to about 380 meters. Thus *>S'. acanthias
(p. 30) occurs in numbers anywhere' between the surface and
the 160-180 meter level. *8. fernandinus ranges from close to
the surface (reported at 16-20 meters by Poll, 1951, p. 62) down
to the maximum depth reported above for its genus, while the
center of abundance lies considerably deeper for it than for
* acanthias, to judge from Poll's (1951, p. 61) report of 608
specimens ti-awled at 220 meters off equatorial West Africa. And
it seems that neither *cuhensis (reported from 137-384 meters)
nor *megalops rise, normally, as near to the surface as does
*acanthias or * fernandinus.

Species. All known representatives of the genus Squalus from
various parts of the world (under whatever names thej^ may have
been reported) seem referable to one or other of three divisions
as defined by the shape of the pectoral fins; by the position
(farther forward or rearward) of the first dorsal fin; by the
presence or absence of a small secondary lobelet on the outer edge
of the flap-like expansion of the anterior margin of the nostril ;

BIQELOW AND SCHROEDER : SQUALOID SHARKS 27

and by the color pattern. In the following discussion each of
these divisions is referred to under the name of its earliest de-
scribed representative.

The differences in the shape of the pectorals, as outlined be-
low, are so obvious that no further discussion of them is called
for just here. Similarh', the position of the first dorsal spine,
relative to that of the origin of the pectoral fins, has proved
definitely alternative for the specimens yet measured, though
with the reservation that measurements of a larger number may
eventually narrow the gap in this respect. And the shape of the
narial flap seems equally diagnostic for the majority of specimens,
though we have seen one individual of the fcrnandinus division
where one of the nostrils shows the secondary lobelet, but the
other does not — a situation analogous to that of the shape of
the inner corner of the pectoral fins in Centrophorus (p. 67).
But our earlier selection (Bigelow and Schroeder 1948, p. 455;
Bigelow, Schroeder and Springer 1953, p. 221) of the midpoint
of the bases of the pelvics relative to the positions of the dorsals
as a species-criterion was a less happy one, for additional in-
formation has shown that in this respect the extremes intergrade.
And while the presence of white spots is diagnostic for the speci-
mens that show them, the absence of white markings is not diag-
nostic, for it has long been known that in the type species of the
genus {acanthias) these spots tend to fade with growth, while
some large specimens show no ti'ace of them.

Synopsis of divisions of Squalus

Acanthias division. Inner corner of pectoral fins well rounded ;
the distal margin not deeply concave; point of emergence from
skin of 1st dorsal spine is posterior to 5th gill openings by a dis-
tance as long as from anterior margin of eye to 5th gill openings
or slightly longer ; midpoint of base of pelvics about midway be-
tween a perpendicular at point of emergence of 2nd dorsal spine
and one at free rear tip of 1st dorsal on most specimens, but
nearer to tip of 1st dorsal than to 2nd dorsal spine on one large
male from Massachusetts (Mus. Comp. Zool. No. 35863) that we
have seen. Anterior narial flap simple, without secondary lobelet
on its outer edge. Sides of trunk marked with white spots on
small and middle sized specimens, more faintly so on larger ones.

28

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Fig. 3. Dermal denticles from side of body below first dorsal fin. A,
Squalus f.ernandirius, female, 914 mm. long, island of Juan Fernandez, Mus.
Comp. Zool. No. 841, x about 50. B, S. fernandirm^, male, 397 mm. long, off
South Carolina, Mus. Comp. Zool. No. 37064, x about 50. C, S. megalops, same
specimen as in Fig. 4A, x about 70. D, Squalus acnnthias, Massachusetts,
X about 34, after Bigelow and Schroeder 1948, Fig. 88C. E, Etmopterus
spinax, female, 250 mm. long, Norway, Mus. Comp. Zool. No. 1022, x about
15. F, Etmopterus princeps, female, 165 mm. long, offing of Nantucket, Mus.
Comp. Zool. No. 37403, x about 12. G, Etmopterus princeps, female, 610
mm. long, offing of southern edge of Georges Bank, Mus. Comp. Zool. No.
37445, X about 10.

BIGELOW AND SCHROEDEB: SQUALOID SHARKS

29

while some adults lack these spots altogether. Dermal denticles
on sides below 1st dorsal fin of the shape shown in Figure 3D.

Fernandinus division. Inner corner of pectorals about a right
angle, more or less blunted, the distal margin weakly concave ;
point of emergence from skin of 1st dorsal spine is posterior to
5th gill openings by a distance at least not longer than from an-
terior edge of eye to 2nd gill openings, and slightly shorter than
that in most ; midpoint of base of pelvics at least as near to a per-
pendicular at rear tip of 1st dorsal as to one at emergence from

A '^

Fig. 4. Outlines of fins of Squalus megalops and of Squalus cubensis,
superimposed to show the differences in shape. A, caudal fins of S. mega-
lops, female 526 mm. long, New South Wales, Mus. Comp. Zool. No. 38619
(dotted line), and of -S. cubensis, female, 672 mm. long, Cuba, Mus. Comp.
Zool. No. 1461 (solid line), adjusted to equal lengths along upper margin.
B, pectoral fins of S. megalops (type specimen), from drawing kindly con-
tributed by Dr. G. P. Whitley (dotted line), and of S. cubensis, same
specimen as in A (solid line), adjusted to equal lengths along outer margin.

skin of 2nd dorsal spine, and nearer on most (but see above,
under acanthias division) ; anterior narial flap with a small sec-
ondary lobelet on its outer edge (Bigelow and Schroeder 1948,
Fig. 87F), though in occasional specimens this lobelet may be
lacking on the one nostril or on the other (see above) ; dermal
denticles on sides below first dorsal fin simpler in form than in

30 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

acanthias division, tridentate, the marginal points longer on the
denticles of small specimens than on those of larger (Fig. 3, A,
B). Trunk without white markings at any stage in growth.

Megalops division. Inner corner of pectoral fin pointed, or
only very slightly blunted, the distal margin deeply concave (Fig.
4B) ; relative positions of fins, shape of narial flap, and color
(without spots) essentially as in fernandinus division (see
above) ; dermal denticles on sides below first dorsal fin differing
from those both of the axanthias division and of the fernandinus
division in a way easier presented pictorially (Fig. 3C) than
verbally.

Remarks. There is no danger of confusing any representative
of the megalops division with an}'- of the acanthias division or any
of the fernandinus division, so sharply diagnostic is the' shape of
the pectoral fins. But the discontinuity as regards fin characters
is so small between the acanthias and the fernandinus divisions
that identification as the one or the other of large plain-colored
specimens may call for close scrutiny of the shape of the narial
flap and of the denticles.

Acanthias division. *Squalus acanthias Linnaeus 1758, type
species of the genus® and locally the most numerous of sharks,
represents this division in the North Atlantic, where its regular
range extends in the east from Iceland, Norway and the Murman
coast to Morocco (including the Mediterranean and the Black
Sea), with reports from the Canaries and Senegal; and in the
west from southeastern Labrador (straying to southwest Green-
land) to the southern part of the North Carolina coast in abun-
dance, with a few ranging to southern Florida and to Cuba.'

The common spiny dogfish of the coast of the northwestern
Pacific was originally described as a separate species, *suckleyi
Girard 1854, and it has most often been reported under that
name. But our own comparison of Californian specimens with
others from both sides of the North Atlantic has not revealed
any consistent differences that might be regarded as specific,
whether in the position of the first dorsal fin relative to the
pectorals (the character the most often invoked as alternative),
or in any other respect (Bigelow and Schroeder 1948, p. 453).

6 For synonyms see Bigelow and Schroeder 1948, p. 467.
" For details, see Bigelow and Schroeder 194S, pp. 463-464.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 31

Spiny dogfishes of the acanthias division have been reported
from northeastern Asia both as *acanthias Linnaeus 1758 and
as *sucMeyi Girard 1854 ; from Japanese waters under the name
vulgaris Risso 1826, repeatedly as *suckleyi Girard 1854, as
mitsukurii Jordan and Fowler 1903 (their illustration, but not
their description, which was based on an example of the fer-
nandinus division, see below, p. 33), and as wakiyae Tanaka
1918, a name proposed by Tanaka for the shark that had been
pictured by Jordan and Fowler (1903, p. 630, fig. 3) as mitsu-
kurii; from Korea, China and Formosa as suckleyi. There is
nothing, however, in available information to suggest that these
populations differ specifically, one from another, or from the
typical * acanthias of the North Atlantic. Schmidt (1931, p. 7)
indeed, has long since expressed a doubt as to whether the spiny
dogfishes of the acanthias division of Japan are separable from
those of the Black Sea. For North Pacific references to Sqvalus
of the acanthias division, see Bigelow and Schroeder 1948,
p. 472.

A representative of the acanthias division has also been cred-
ited (by name only) to Hawaii (Fowler 1930, p. 495, as S.
suckleyi). But an earlier report of mitsitkurii from Hawaiian
waters proves actually to have been based on *S. fernandinus
(p. 33).

The acanthias division has not been reliably reported for the
equatorial-subequatorial belt, whether of the Atlantic or of the
Indo-Pacific. But seemingly it is as widespread in mid-latitudes
of the southern hemisphere as it is of the northern. Thus sharks
apparently of this division have been reported from southern
Africa as *acanthias and as vulgaris ; from Uruguay and north-
ern Argentina as ^acanthias; from the Magellanic region as
"^acanthias and as lehruni Vaillant 1888^ ; from Chile as ^suckleyi
(by Fowler 1930, p. 495) ; from New Zealand as * fernandinus
(by Waite 1909, p. 142, PL 16, fig. 1) but identity as acanthias-
like established by rearward position of first dorsal fin and by
white spotted sides, and as kirki Phillipps 1931 ; from Australia
as vulgaris (repeatedly) and as whitleyi Phillipps 1931 (see also
\Vhitley 1940, p. 193); and from the island of Reunion in the
southern Indian Ocean as vulgaris. No one of these southern
populations, however, has yet been studied in detail, nor has any-
thing developed within the past few years to contradict our

32 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

earlier statement (Bigelow and Schroeder 1948, p. 454) that "the
relationsliip of these southern hemisphere forms to one another
and to the northern acanthias is uncertain."

Fernandinus division. The earliest named representative of
this division is Squalus fernandinus Molina 1782, originally
described from the island of Juan Fernandez off the coast of
Chile. And our recent comparison of a female, 520 mm. long
(Mus. Comp, Zool. No. 446) from Nice, France, with a * fernan-
dinus of 914 mm. from the type locality, as well as with others
from the Gulf of Mexico and from the offing of South Carolina,
has confirmed the conclusion earlier arrived at by Garman (1913,
p. 195), by Poll (1951, p. 59), and by Bigelow, Schroeder and
Springer (1953, p. 222), that the shark described from the Medi-
terranean in 1826 by Risso under the species-name hlainvillii
(Risso 1826, p. 133, PI. 3, fig. 6) is identical with fernandimis.
This joint species (most often reported as ^^Nainvillii^^) occurs
widespread in the Mediterranean, in the Black Sea, and off the
open Atlantic coast from Portugal southward to equatorial
waters (reported from 6° 31' S., see Poll 1951, p. 59). And it is
to be expected all along the West African seaboard to the south-
ward, for spiny dogfishes apparently identical with * fernandinus
have been reported in abundance in southern African waters,
both as hlainvilUi (by Bleeker, 1860, pp. 50, 58, 80, and by Giin-
ther 1870, p. 419) : as acutipinnis Regan 1908 (by Regan 1908\
p. 248, and by Barnard 1925, p. 48), and as * fernandinus (by
Gilchrist 1922, p. 48, by von Bonde 1924, p. 5, and by Smith 1949,
p. 60). *;S'. fernandinus is known in the western side of the At-
lantic also, both from the offing of South Carolina and in the Gulf
of Mexico in the north (Bigelow, Schroeder and Springer 1953,
p. 220), and off the coast of Argentina in the south (for locality
records from the western South Atlantic, see Bigelow and Schroe-
der 1948, p. 480). But the number of specimens so far reported
from the western side of the Atlantic, north or south, has not
been large enough to suggest that *fer7iandinus is as plentiful
there, or as generally distributed, as it is in the eastern side, and
in the Mediterranean.

The reports that have come to hand for *fer7iandinus in the
Indo-Pacific are numerous enough, and are distributed widely
enough to show that it is as widespread there as it is in the At-
lantic, not only in the temperate zones both north and south but

BIGELOW AND SCHROEDER : SQUALOID SHARKS 33

in tropical-subtropical latitudes as well. Thus *fernandinus, or
a form so closely allied to it that no significant differences appear
from the published accounts, is widespread in Japanese waters,
whence it has been reported as mitsukurii Jordan and Fowler
1903 (their description, but not their illustration, which was
based on a specimen of the acanthias division, see above p. 31^) ;
repeatedly also as japonicus Tanaka 1917 (for an excellent de-
scription and illustration, see Tanaka 1917, p. 467, PI, 30, figs.
366, 367). It has also been described and pictured from southern
China by Fang and Wang (1932, pp. 249, 250, fig. 16), not
only under the name mitsukurii Jordan and Fowler 1903, but
also as brevirostris Tanaka 1917, though this last name had
actually been based on the Japanese representatives of the meg-
alops division, not on one of the fernandinus division (see below,
p. 36).

The specific identity of the spiny dogfishes that had been re-
ported by Snyder (1904, p. 515), by Jordan and Evermann
(1905, p. 45), by Gilbert (1905, p. 580), and by Giinther (1910,
p. 490), from the Hawaiian Islands under the name mitsukurii
Jordan and Fowler 1903, had remained uncertain, for the brief
descriptions seemed equally applicable to the fernandinus di-
vision of the genus or to the megalops division. But our recent
examination of five of Jordan and Evermann 's and Gilbert's
specimens has shown nothing to separate them specifically from
the specimens of ^fernandinus which we have seen from the offing
of South Carolina, from the Gulf of Mexico, from the Mediter-
ranean, and from Juan Fernandez (type locality), whether in
relative positions of fins, in bodily proportions, or in the shape of
the denticles on the sides of the body.

*S. fernandinus has also been reported from the Philippines
under its own name (with brief description) by Herre (1923, p.
73; 1934, p. 12) ; also as philippinus Smith and Radcliffe (1912,
p. 677, PI. 51, fig. 1), a supposedly new species based, however,
on a specimen that is not separable from fernandinus either in
fin characters or in its denticles, as Mr. Stewart Springer informs
us from a recent examination of it in the U. S. National Museum.
We should note in passing that Whitley (1931, p. 310) had pro-
posed a new name, montalbani, to replace philippinus Smith and
Radcliffe, the latter name having long been preoccupied within

8 On this point, see also Jordan and Hubbs 1925, pp. 105-106.

34

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

a

S3

o

BIOELOW AND SCHROEDER : SQUALOID SHARKS 35

the genus Squalus by philippimis Shaw 1804 (p. 341), proposed
for an Australian heterodontid. For details see Whitley.

*S. fernandinus, to continue, has been reported from Mauritius,
both as hlainvillei (by Sauvage 1891, p. 11) and as acutipinnis
(by Regan 1908, p. 47). It has long been known also in Aus-
tralian waters, where it seems first to have been reported, with
at least presumptive evidence of identity, by Giinther (1870, p.
419) as 'blainvillei, and subsequently by that same name but with-
out description by MacLeay (1881), by Ogilby (1889, p. 185),
and by Lucas (1890, p. 44). A confusing feature here is that the
only Australian report of a spiny dogfish under the name *fer-
nandinus, the identification of which was supported by a de-
scription or by an illustration (Waite 1921, p. 23, fig. 30), seems
not to have been based on that species, but on the local repre-
sentative of the megalops division of the genus.

Eegan's (1908, p. 46) identification, however, of a spiny dog-
fish from Tasmania as * fernandinus was no doubt correct, for we
have found nothing to separate the type specimen of *tasmanien-
sis Rivero 1936 (now in the Museum of Comparative Zoology)
from our specimen of *fernandinus from the type locality of the
species (Juan Fernandez). For an extensive list of Indo-Pacific
records for fernandinus see Herre 1953, p. 31.

With the evidence of widespread occurrence so conclusive for
fernandinus for the Australian region in general, it seems
astonishing that there has been no report yet for New Zealand
waters of a spiny dogfish, the accompanying account of which
places it definitely in *' fernandinus. In fact, the only report under
that name of a New Zealand Squalus that includes any clear evi-

Fig. 5, A, Etmopterus bnllisi, new species, juvenile male, 230 mm. long
(U. S. Nat. Mus. No. 158186), off northeastern Florida. B, upper teeth of
same, x about 9. C, lower teeth of same x about 12. D, dermal denticles
of same, from side below first dorsal fin, x about 30. E, Etmopterus polli,
immature male about 226 mm. long, off south equatorial West Africa, Mus.
Comp. Zool. No. 38000, lower surface of anterior part of head, x about 1.4.
F, Etmopterus spvnax, Norway, to show black pattern on posterior part of
trunk. G, corresponding drawings of E. virens, northern part of Gulf of
Mexico. H, corresponding drawings of E. hillianus, north coast of Cuba;
F-H, X about 0.4, after Bigelow, Schroeder and Springer 1953, Fig. 6. I,
corresponding drawing of E. ahernathyi, type specimen, x about 0.4, after
Garrick 1957, Fig. 3C.

36 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

dence of specific identity (Waite 1909, p. 142, PI. 16, fig. 1;
Phillipps 1928, p. 223, fig. 3) was evidently based on a member
of the acanthias division. Thus the accompanying illustration
shows its sides as white spotted, its first dorsal fin as originating
posterior to the rear inner corner of the pectorals when the lat-
ter are laid back, and the mid-point of the bases of its pelvics
as nearer to the origin of the second dorsal than to the rear end
of the base of the first dorsal.

It is astonishing, also, that while *fernandinus must be com-
mon around Juan Fernandez, to judge from the number of speci-
mens that have found their way thence to various museums (Del-
fin 1901, p. 21), and is reported from northern Argentine waters,
on the conclusive evidence of a small one found in the stomach
of an albatross (Lahille 1928, p. 327, fig. 17), we have found no
positive report of its presence in continental waters along the
coast of Chile. Thus Philippi's (1887, p. 27) inclusion of it
(under the name firmandezianus) in his account of the sharks
of Chile seeminglj^ harked back to Molina's original report of it
from Juan Fernandez. Neither does Delfin (1901) mention any
Chilean localities for it other than Juan Fernandez, which is
some 240 miles ofl^ the Chilean coastline ; nor does Fowler's (1930,
p. 495) nominal reference of it to "Chile" include any con-
tinental locality. Neither has it been reported from Peru, nor
from the Pacific coast of Central or North America, for that
matter.

Megalops division. The earliest adequate account (with illus-
tration) of a representative of this was by McCulloch (1927, p.
9, PI. 2, fig. 25a; illustration later copied by "Whitley 1940, p.
138, fig. 147) of an Australian shark that was identified by him
as *megalo2)s MacLeay 1881. And Dr. Whitley writes us that he
has verified this identification, by comparison with the type
specimen of *megalops, which he has had the kindness to ex-
amine at our request. The accompanying illustration (Fig. 4B)
reproduces a tracing of its pectoral fin which Dr. Whitley has
kindly sent us. We are also fortunate enough to have at hand
three excellent specimens of *megalops, 526-550 mm. long, from
New South Wales, for which the Museum is indebted to Col.
John K. Howard.

The Japanese brevirostris, described and beautifully pictured
by Tanaka (1917, p. 464, PI. 129, figs. 362, 363; PI. 130, fig. 364)

BIQELOW AND SCHROEDER : SQUALOID SHARKS 37

resembles *m€lagops so closely that we have no doubt a com-
parison of specimens from the two regions would lead to their
union as ^megalops, this being the older of the two names.

The *megalops division is represented in the Atlantic by
*cuhensis Rivero 1936, which is known from Cuban waters (type
region), from the northwestern part of the Gulf of Mexico,
(specimen recently taken by "Oregon" and referred to us by
Mr. Stewart Springer), probably Trinidad and from Rio de Jan-
eiro, whence it was reported by Ribeiro (1907, p. 168) under the
name hlainvillei (Bigelow and Schroeder 1948, p. 478, footnote
63).

*S. cuhensis closelj^ resembles *;S^. megalops in proportional
dimensions, and in the shape of the denticles on the sides of its
body (Fig. 3), but differs in a relatively longer first dorsal fin
spine, in somewhat differently shaped caudal and 'pectoral fins
(Fig. 4), and in the location (relative to the dorsal fins) of the
pelvic fins, the midpoint of the bases of which are about midway
between the rear end of the base of the first dorsal and the
origin of the second dorsal in * cuhensis (a little nearer to the
rear end of the base of the first dorsal than to the origin of the
second dorsal in * megalops). No representative of this division
of the genus has yet been reported from the eastern side of the
Atlantic, or from the Mediterranean.

Genus CiRRHIGALEUS Tanaka 1912

Cirrhigaleus Tanaka 1912, type species C. harbifer Tanaka 1912b, p. 151,

PI. 41, figs. 156-162; Japan.
Generic synonym :

Phaenopogon Herre 1935, type species P. harhulifer Herre 1935, p.

123, fig. 1; Japan.
Generic characters. In general as in Squalus (p. 26), but
without precaudal pit either above or below; posterior part of
trunk without longitudinal ridges; and anterior margin of nos-
tril, near inner end, with a well developed and very conspicuous
fleshy barbel tapering to a slender tip (a feature unique among
squaloid sharks) ; dermal denticles on sides of body with tri-
dentate rear margins, their outer surface with three correspond-
ing ridges. Also, in the only species that has yet been seen, the
back is more highly arched than it is in any known Squalus, and

38 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the body is subtriangular in cross section. Maximum recorded
length 855 mm.

Remarks. Garman (1913, p. 457), followed by Fowler (1941,
pp. 255, 275) reduced Cirrhigaleus to a subgenus of Squalus in
rank. But the presence of the nasal barbel, combined with the
lack of precaudal pits and of longitudinal-lateral ridges on the
caudal peduncle seem fully to justify its retention as a distinct
genus. Phaenopogon Herre, proposed without mention of Tan-
aka's (1912) earlier account of Cirrhigaleus, is clearly equivalent
to the latter, to the synonymy of which Herre himself (1936,
p. 59) has already relegated it.

Species. One species only is known, C. harhifer Tanaka 1912,
which was based on a single male, 855 mm. long, from the Tokyo
market. A second specimen (a female of 555 mm.) from Misaki,
Japan, has more recently been described and pictured by Herre
(1935, p. 123, fig. 1) as Phaenopogon barhulifer.

Genus CeNTEOSCYLLIUM Miiller and Henle 1841

Centroscyllium Miiller and Henle 1841, p. 191. Type species *Spinax fabricii

Eeinhardt 1825, West Greenland.
Generic synonym:

Paracentroscyllium Alcock 1889, p. 379, type species P. ornatum Alcock,

Bay of Bengal.
Generic characters. Squalinae with upper teeth similar to
lowers, erect and symmetrical, or nearly so, all along the jaw ;
the teeth in both jaws with 3-5 cusps, the median cusp much
the largest; dorsal fin spines well developed, with two longi-
tudinal grooves on either side ; lower posterior margin of caudal
fin with subterminal notch (but see discussion, p. 46) ; caudal
peduncle without precaudal notch either above or below, and
without lateral-longitudinal ridges ; inner corner of pectoral fins
rounded in known species; snout in front of mouth not more
than one-half as long as head to origin of pectoral fins; pre-
oral clefts present, slightly expanded inwardly; dermal denti-
cles thorn-like, on stellate bases; newborn specimens naked, the
denticles developing first along the back, spreading later to sides,
and subsequently to the lower surface (see further, p. 41) ;
some species, at least, with thickenings of the skin that presum-
ably are luminous. Maximum recorded length 829 mm., for C.
fahricii.

BIQELOW AND SCHROEDER : SQUALOID SHARKS 39

Depth range. The only member of this genus for which we
have more than scattered records {fabricii, p. 41) seldom (if
ever) has been taken shoaler than 275 meters; its center of
abundance lies deeper than about 500 meters. It is known to
occur down at least to 1100 meters, and there is a somewhat
questionable report of it from about 1495 meters (for details
see Bigelow and Schroeder, 1948, p. 485). Depth records for
other members of the genus have been from between 448 and
1024 meters.

Remarks. The few known species that fall in Centroscyllium
as generally defined are set apart from all other known mem-
bers of the subfamily Squalinae by their dentition, the teeth
having 3-5 cusps in both jaws. In this respect they are ap-
proached the most nearly by Etmopterus, in which the uppers
have 3-7 cusps, but the lowers have only one. In all other known
members of the subfamily the teeth (uppers as well as lowers)
have only one cusp.

The genus Paracentroscyllium was proposed by Alcock (1889,
p. 379) for three juvenile sharks, 5i/4 inches long, from the Bay
of Bengal, allied to Centroscyllium except that they had "mono-
cuspid teeth" and an "absolutely smooth integument." But the
nakedness of the skin was clearly a juvenile character (this
accords with conditions in *fahricii, see p. 41) ; and the lateral
cusps on the teeth seem merely to have escaped notice, for Al-
cock's (1896, p. 308) subsequent study of four additional speci-
mens up to nearly one foot long from the Arabian Sea led him
to emend his original diagnosis of ornatum to read "minute tri-
cuspid teeth in both jaws" and body "covered with minute
placoid deciduous scales." Accordingly, in 1896 (pp. 308, 310)
he united Paracevitroscyllium with Centroscyllium. Hence it
was as a Centroscyllium that he pictured ornatum in 1900 (PI.
35, figs. 1-lB, including a figure of one of its denticles). And
Fowler's (1941, p. 252) revival of Paracentroscyllium, as a sub-
genus of Centroscyllium, for species in which the "caudal ends
in point posteriorly, lower edge of fin without distinct posterior
notch," (harking back, no doubt, to Regan's [1908, p. 40] key
to the species of Centroscyllium), runs counter to Burkhardt's
(1900, p. 567, fig. 8) illustration of one of Alcock 's original
specimens, for this shows the caudal as blunt-tipped and with
a distinct (though shallow) subterminal notch.

40

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

CO

OS

o

BIGELOW AND SCHROEDER : SQUALOID SHARKS 41

Species. The following supposedly separate species fall in
Centroscyllium : *fabricii Reinhardt 1825 (type species), both
sides of the northern Atlantic ; grayiulatum Giinther 1887, Falk-
land Islands ; ornatum Alcock 1889, Bay of Bengal and Arabian
Gulf; *nigrum Garman 1899 (inel. *ruscosiim Gilbert 1905),
eastern Pacific off Panama (vicinity of the Galapagos, vicinity of
Cocos Island), also Hawaiian region; and *'ritteri Jordan and
Fowler 1903, Japan.

*C. fahricii, the common ''black dogfish" of English-speaking
North Atlantic fishermen, has been described by several authors.
Even so, examination of large catches made recently off the Nova
Scotian coast (Bigelow and Schroeder 1954, p. 40) enables us
to expand our earlier account (Bigelow and Schroeder 1948, p.
482) in the following respects. The teeth are already tricuspid
in both jaws on specimens not more than 165 mm. long (suggest-
ing that this is already the case at the time of birth), and the
skin, then, is wholly naked above as well as below, just as Alcock
(1889, p. 379) described it for the original (133 mm.) speci-
mens of his ornatum, from the Bay of Bengal. But a * fahricii of
171 mm. already shows a dense band of denticles along the back,
with a few on the upper part of the sides below the second dor-
sal fin. On one of 177 mm., the back as a whole, and the sides
above the mid-level are rough with denticles, though the lower
surface still is smooth. And the situation is much the same on
one of 273 mm. But on specimens of 300 mm. and larger, the
belly, like the upper parts, is strewn with denticles.

The color pattern, too, undergoes an interesting alteration
with growth. The smallest specimen we have seen (165 mm.
long) is ink-black below and slightly paler along the back and
on the basal parts of the fins, with the two dorsals and the
pectorals margined and the pelvics tipped with white, and with

Fig. 6. A, Etmopterus lucifer, female 337 mm. long, Japan, Mus. Comp.
Zool. No. 1109. B, outlines of juvenile male, 267 mm. long, Japan, Mus.
Comp. Zool. No. 1123, to show the distribution of the black, presumably
luminescent, markings. C, denticles from side of body below first dorsal
fin of specimen shown in A, x about 35. D, Etmopterus brachyurus, male,
300 mm. long, Japan, after Smitii and Radcliffe with the black markings
added from the type specimen in the U. S. National Museum. E, ventral
view of posterior part of trunk of same specimen.

42 BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY

white fin spines. The development of the denticles (see above)
is accompanied by a paling to deep chocolate brown, first along
the mid-zone of the back, then along the upper part of the sides,
as illustrated by specimens of 171-177mm. (see above). The
belly, however, continues black and the fins continue white-
edged up to a length of 200-225 mm. But it is only the extreme
tips of the dorsal fins that show any trace of white by the time
a length of 280-330 mm. is reached, and all the fins on larger
specimens are uniformly blackish right out to the margins.
Meantime, the back and the upper part of the sides darken again
after the dermal denticles have spread to the lower surface, so
that half-grown specimens, and larger, are black above as well as
below when taken from the water, unless the skin has been
rubbed off, as often happens in the trawl. We have seen no ex-
ception to this rule among the many specimens we have handled
during the cruises of "Caryn" and of "Capn. Bill II" during
1949, 1952 and 1953 (Bigelow and Schroeder 1954, p. 41). Our
earlier account of the color of adult *fahricii as "deep chocolate
brown, darkest (almost black) below and on the fins generally"
(Bigelow and Schroeder 1948, p. 485) was based on a specimen
that had reposed for several years in alcohol.

We think it evident, from the foregoing, that changes with
growth must be taken into account in any discussion of the pos-
sible taxonomic significance of fin-markings and of the regional
distribution of dermal denticles in this particular genus.

The North Atlantic, Japanese, and Falkland Island repre-
sentatives of the genus all seem to deserve continued recognition
as separate species, though they resemble one another closely in
general appearance. Thus the distance from the upper origin of
the caudal fin to the rear end of the base of the second dorsal
fin is as long as from the eye to the level of the 4th-5th gill
openings in *ritteri of which we have an excellent specimen at
hand, about 425 mm. long (Fig. 7A), whereas in *fahricii of
about that same size the caudal peduncle (measured as above) is
only about as long as from the eye to the first gill openings. This
contrast, indeed, was invoked as alternative between the Japan-
ese and North Atlantic populations in our earlier key to the
species of Centroscylliuni (Bigelow and Schroeder 1948, p. 482).
The denticles, too, on specimens of about equal sizes are some-
what sparser on *ritteri than on *fahricii; as an illustration,

BIGELOW AND SCHROEDER : SQUALOID SHARKS 43

7 or 8 denticles on the side, below the first dorsal fin, counted
along a nearly straight line, occupy a length of 1 cm. on our
*ritteri of 425 mm., but 8 to 12 denticles occupy this space on
the specimens of *fahricii of about 400 mm.

The general coloration of the body seems also to be distinctive
here, for all the *fahricii we have handled, large enough for
the denticles to have spread to the lower surface (p. 41), have
been plain above as well as below, not only when fresh but after
a few years' stay in alcohol. But the two original specimens of
*ritteri, taken three years previous, are described as "uniform
dark grayish brown, blackish below in front" (Jordan and
Fowler, 1903, p. 635). And this applies equally to the alcoholic
specimen of ^ritteri we have at hand, the lower surface of which
is densely- denticulate, while it also shows a definite black flank
mark on either side of the rearward part of the trunk (Fig. 7A).

Judging from Burkhardt's (1900, p. 567, fig. 7) outline draw-
ing of the type specimen (270 mm. long), C. granulatum
Giinther 1887, from the Falkland Islands, agrees more closely
with *ritteri than with *fabricii in the length of the caudal
peduncle relative to the dimensions of the head. But Giinther 's
(1887, p. 7) characterization of the dermal denticles of granu-
latum as "much coarser" than those of '*fahricii "and in the
form of granulations" sets it apart equally from *ritteri, the
thorn-like denticles of which (Fig. 7C) more nearly recall those
of *fabricii (Bigelow and Schroeder 1948, fig. 91), except that
thev are relativelv somewhat longer and more slender.

With *riiteri clearly separable from *fahricii, and granulatum
seemingly so, the same might have been expected to hold good
for ^nigrum (including *ruscosum Gilbert 1905) ; or at the least,
a closer relationship might have been expected with *ritteri than
with *fahricii on geographic grounds. Actually, however, *ni-
grum is pictured both by Garman (1899, PI. 1, fig. 2) and by
Beebe and Tee Van (1941, p. 120, fig. 32), as falling with *fah-
ricii in the shortness of its caudal peduncle relative to the di-
mensions of the head, which w^e can corroborate from our exam-
ination of one of Gilbert's original Hawaiian specimens of his
*ruscosum, about 400 mm. long (the co-type), loaned us by
the Stanford University Museum through the kind offices of Dr.

44

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

H

O

I— I

BIGELOW AND SCHROEDER : SQUALOID SHARKS 45

George Myers,® and of the type specimen in the U. S. National
Museum. Nor can we find anything in the dermal denticles to
separate *nignim from *fahricii.

Also, Garman's (1913, p. 230) use of the number of cusps
on the lower teeth as alternative between * nigrum (with five)
and *fahricii (with three) has failed to withstand the test of
time, for most of the lowers midway along either side of the jaw
actually have five cusps in *fahricii (Bigelow and Schroeder
1948, p. 483, fig. 91E). And Beebe and Tee Van (1941, p. 121)
have found that on their Cocos Island specimens of *nigrum the
outermost pair of cusps were not as prominent as pictured by
Garman (1899, PI. 4, fig. 6).

*C. nigrum, however, maturing at a length of only about 400
mm. (Gilbert 1905, p. 581) or perhaps smaller still (Beebe and
Tee Van 1941, p. 120) is a much smaller shark than *fabricii,
the males of which seemingly do not mature until at least 600
mm. long. Also, the pectoral and dorsal fins continue white
tipped to maturity in *nigrum (Gilbert 1905, p. 581, fig. 230;
Beebe and Tee Van 1941, p. 120, fig. 32) though with the white
areas somew^hat reduced, whereas half-grown specimens of */a&-
ricii, and larger, show no trace of the white fin-markings that
are so conspicuous on small specimens (p. 42). It may also
prove (when more specimens have been examined) that it is
characteristic for the pectorals to extend somewhat farther rear-
ward in *nigrum than in *fabricii, for they are pictured both by

9 The larger of Garman's original two specimens of *nigrum, used as the
basis for his account of the anatomy of the genus GentroscylUum, is now in a
verj- fragmentary condition.

Fig. 7. A, Centroscyllium ritteri, female about 425 mm. long, Mus. Comp.
Zool. No. 1370. B, second dorsal fin spine of same drawn to larger scale to
show the longitudinal grooving. C, dermal denticles from side below first
dorsal fin, x about 6 to 8. D, upper and lower teeth from central part of
jaw, X about 4. E, upper and lower teeth from midway along side of jaw,
X about 4. F, Centrophorus foUaceus, immature male, 351 mm. long, in
U. S. Nat, Mus., from Japan. G, upper and lower teeth of same at center
of mouth, X about 4. H, denticles of same from side below first dorsal fin,
X about 13. I, lateral view of denticle, x about 18. J, Centrophorus
armatus, southern Africa, second dorsal spine of specimen 330 mm, long,
X about 3,5, from a photograph kindly contributed by Dr. J, L. B, Smith
(see p. 6).

46 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Garman (1899, PI. 1, fig. 2) and by Beebe and Tee Van (1941,
p. 120, fig. 32) as reaching nearly or quite to a perpendicular at
the point of emergence (from the skin) of the first dorsal spine,
which is true also of the Hawaiian specimen we have seen (men-
tioned above). It therefore seems justifiable to accept ^nigrum
as a distinct species, at least until a larger number of specimens
from the Pacific populations have been studied with a critical
eye.

C. ornatum of the northern Indian Ocean falls with the fah-
ricii-nigrum-r Uteri group in its thorn-like denticles (Alcock
1900, PI. 35, fig. 1). It agrees more nearly in proportional di-
mensions with *nigrum of low latitudes in the eastern Pacific
than with *ritteri of the northwestern Pacific, the distance from
the rear end of the base of its second dorsal fin to the origin of
the upper side of its caudal fin being only about as long as from
the eye to the first pair of gill openings, while its pectorals reach
rearward about to a perpendicular at the point of emergence of
the first dorsal fin spine. But Alcock (1896, p. 308) describes
the fins as well as the trunk as "uniform jet black" on speci-
mens nearly one foot long. Neither does his account of the type
specimen, 133 mm. long, nor his illustration of it (Alcock 1894,
PI. 8, fig. 2) suggest any white fin-markings, such as are con-
spicuous on nigrum to maturity. While Regan (1908, pp. 40-41)
characterized ornatum as without a distinct posterior notch on
the caudal fin, seemingly following Alcock 's (1894, PI. 8, fig. 2)
original illustration, Burkhardt's (1900, p. 567, fig. 6) drawing
of one of the original series in the British Museum clearly indi-
cates a subterminal notch. And Alcock 's (1900, PI. 35, fig. 1)
second representation of the caudal of a larger specimen seems
more fanciful than realistic, so widely does it differ from his
earlier illustrations. With our knowledge of ornatum so far from
adequate, and with the specimens of it in the collection of the
Zoological Survey of India in a bad state of preservation (the
late Dr. S. L. Hora has so written us in a recent letter), a final
decision as to its status obviously is a matter for some future
student. Meanwhile we believe a tentative key to the species of
Centroscyllium, using the length of the caudal peduncle as the
first alternative, seems likely to be more helpful for identifica-
tion than our earlier key, in which the primary alternative is the
location of the rear margin of the pectoral fin (when laid back)
relative to the location of the first dorsal fin spine.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 47

Provisional Key to Species of C entroscylUum

1. Distance from rear end of base of second dorsal fin to origin of upper
side of caudal fin nearly or quite as long as from eye to level of origin
of pectoral fins *ritteri Jordan and Fowler 1903

Japan (Fig. 7 A,B,C). p. 42
Distance from rear end of base of second dorsal fin to origin of upper
side of caudal fin only about as long as from eye to first-second gill
openings 2

2. Dermal denticles on anterior part of trunk "in the form of granula-
tions" (Giinther 1887, p. 7) fjranuJatum Giinther 1887

Falkland Islands, p. 43
Dermal denticles in the form of spines or thorns on stellate bases .... 3

3. Margin of pectorals, when laid back, falls short of a perpendicular at
point of emergence from skin of first dorsal spine by a distance at
least 25 per cent as long as eye *fabricii Eeinhardt 1825.

North Atlantic, p. 41
Margin of pectorals, when laid back, reaches nearly or quite to below
point of emergence from skin of first dorsal spine 4

4. Dorsal, pectoral and pelvic fins white edged at all stages in growth .

*nigrum Garman 1899
(incl. ruscosum Gilbert 1905), east-
ern tropical Pacific off Central
America, also Hawaiian region, p.
43.

Fins, as well as body, black at all stages in growth

ornatum Alcock 1889
Bay of Bengal and Arabian Gulf. p. 46

Genus EtmOPTEEUS Eafinesque 1810

Etmoptents Eafinesque 1810, p. 14; type species E. aculeatus Eafinesque,
Mediterranean, equals *Squalus spinax Linnaeus 1758. Generic syn-
onyms, see Bigelow and Schroeder 1948, p. 487.

Generic characters. Essentially as in CentroscylUum (p. 38),
except with lower teeth much as in Squalus, i.e., much broader
than the uppers and with only one cusp, the base of each tooth
overlapping that of the next outward so that each row forms a
continuous dental band, and with the cusps directed so strongly
outward all along the jaw that their successive inner margins
act as a nearly unbroken cutting edge (Bigelow, Schroeder and
Springer 1953, figs. 7-10) ; dermal denticles on sides of body

48 BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY

ranging from thorn-like to bristle-like and to truncate, on quad-
rate bases.

Size. At maturity the various species of Etmopterus range
in length from slightly more than 200 mm. {*virens, perhaps also
*villosus) to a recorded maximum of 728 mm. in the case of
*princeps (Collett 1904, p. 4).

Depth range. The members of this genus are creatures of at
least moderately deep water, where (as a group) their depth
range is a wide one. The shoalest captures we find recorded for
any of them are 73-91 meters for *spi7iax in the northernmost
parts of its range (Smitt 1895, p. 1165) and 183 meters for
^abernathyi, off New Zealand (Garrick 1957, p. 181) ; the deep-
est are 2074 meters for E. princeps (see Bigelow, Schroeder and
Springer 1953, p. 251; Bigelow and Schroeder 1954, p. 47; and
Grey 1956, p. 95 for details as to * princeps). Most of the records
for one species or another in various parts of the world have
been from about 300 meters to about 900-1000 meters.

Remarks. These small, black-bellied deep-water sharks, of
which some 16 recognizably distinct species are known, form a
very compact unit. Characters that have proved the most use-
ful for diagnosis of species within the genus are : the sizes and
relative positions of the fins; the color pattern, whether uni-
formly as dark above as below or paler above with conspicuous
dark flank markings of characteristic shape, whether or not with
complex patterns of longitudinal bands of black dashes and dots
(presumably luminescent) and whether with or not with white
fin markings; also, for half-grown specimens and larger, the
shapes of the dermal denticles, and their arrangement. But in
using the denticles for diagnosis allowance must be made for the
size of the specimens in question, for we have found that in one
species {"^'princeps) where the denticles are bristle-like on very
young specimens, they are succeeded by thorn-like forms on half-
grown individuals and larger (Fig. 3, F, G).

For accounts of the luminescent organs (for the species that
have them) and of luminosity in this genus, see Johann (1899)
and Oshima (1911).

Whitley (1939, p. 266) has revived the genus Acanthidium
Lowe 1839, type species *A. pusillum Lowe 1839 {=Centrina
nigra Lowe 1834, preoccupied), designated by Jordan and Ever-
mann (1896, p. 55), as distinct from Etmopterus Rafinesque

BIGELOW AND SCHROEDER : SQUALOID SHARKS 49

1810, type species E. aculeatus Rafinesque, {=*Squalus spinax
Linnaeus 1758). And Fowler (1941, p. 251) has recognized
two corresponding subgenera within the genus Etmopterus,
namely Acanthidivm Lowe 1839, with "second dorsal origin be-
hind ventral base ; first dorsal origin midway between orbit and
second dorsal origin," and Etmopterus Rafinesque 1810 with
"second dorsal origin over or a little before ventral base; ventral
origin slightly nearer subcaudal origin than pectoral origin."
But these differences do not seem to us wide enough for generic
— or even subgenerie — separation (p. 22; Bigelow, Schroeder
and Springer 1953, p. 238). Acanthidium as employed by Gar-
man (1913, p. 215) in a different sense, is discussed on p. 21.

Species. The genus Etmopterus is especially interesting from
the taxonomic standpoint, because it is already known to include
a larger number of recognizable species than we have any reason
to suppose is included in any other genus of squaloid sharks.
These species, it is true, resemble one another so closely in gen-
eral aspect that any one of them might easily be mistaken for
any other, on cursory inspection. But closer examination has
shown that they are sharply separated by differences both in the
relative positions of the fins and in the nature and arrangement
of the dermal denticles, and also in the color pattern which has
proved a much more convenient species-character here than it is
usually thought to be among other sharks.

The following seven species have been known from the North
and Equatorial Atlantic, including the Mediterranean and the
Gulf of Mexico.

1. *spinax Linnaeus 1758, type species of the genus (many of
the reports of it have been as niger Cloquet 1820, p. 93) ; eastern
North Atlantic (including the Mediterranean) from the Cape
Verde Islands, Morocco and the Azores to Norway.

2. *pusillus Lowe 1839 {=Centrina nigra Lowe 1834, but
nig7'a is preoccupied in Etmopterus by niger Cloquet 1820, pro-
posed in substitution for spinax Linnaeus 1758 and employed by
many subsequent authors) ; tropical-subtropical Atlantic, from
tropical West Africa to the Canaries, Madeira, and the Azores
in the east (including the Mediterranean) and Gulf of Mexico
in the west, where it has been found recently (Bigelow,
Schroeder and Springer 1955, p. 2). The Japanese shark de-
scribed and pictured under the name pusillus by Tanaka (1912,

50 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

PI. 22; 1912^, p. 88) was based on a clearly separable form (see
below, p. 58).

3. *hillianus Poey 1861, Cuban-West Indian region, north-
ward to the ofiEing of Chesapeake Bay.

4. *princeps Collett 1904, both sides of the North Atlantic,
P'aroes-Hebrides region and offing of Gibraltar in the east; con-
tinental slope in the west where it is common from the offing of
southern Nova Scotia to that of southern New England (Bigelow
and Schroeder 1954, p. 46).

5. *polli Bigelow, Schroeder and Springer 1953, equatorial
West Africa.

6. *schultzi Bigelow, Schroeder and Springer 1953, northern
part of the Gulf of Mexico.

7. *'virens Bigelow, Schroeder and Springer 1953, northern
part of the Gulf of Mexico.

For the diagnostic characters of the members of this group,
see Bigelow, Schroeder and Springer 1953, p. 237.

An eighth North Atlantic Etmopterus, from the east coast of
Florida, is described here as a new species, *E. hnllisi.

Etmopterus bullisi n. sp.

Figure 5 A-D

Plate 2

Type specimen. Female, 196 mm. long; "Pelican" Sta. 42, off
northeast coast of Florida, Lat. 30°02' N, Long. 80°05' W; 205
fathoms (U. S. Nat. Mus. No. 158186).

Additional material : female of 205 mm. and immature male of
230 mm. from this same locality; also immature male of 212
mm., "Pelican" Sta. 51, off eastern Florida, Lat. 29°48' N, Long.
80°09' W, about 200 fathoms.

Description. Proportional dimensions in per cent of total
length of type specimen, and of a male 230 mm. long, to nearest
0.1 per cent.

Snout length in front of: outer nostrils 2.5, 2.8; mouth 11.7, 12.2.

Eye: horizontal diameter 4.6, 4.6.

Mouth: breadth 7.1, 7.0.

Nostrils: distance between inner ends 2.8, 2.8.

Gill openings: lengths first 2.3, 1.7; third 2.3, 1.7; fifth 2.0, 1.5.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 51

First dorsal fin: vertical height 3.1, 2.8; length of base 4.6, 4.4.

Second dorsal fin: vertical height 4.6, 4.4; length of base 7.4, 7.0

Caudal fin: upper margin 24.0, 23.5.

Pectoral fin: outer margin 11.0, 10.0; inner margin 4.8, 5.6, width 4.6, 5.2.

Distance from snout to: 1st dorsal 33.7, 33.0; 2nd dorsal 56.6, 57.9;
upper caudal 76.0, 76.5; pectorals 25.0, 24.4; pelvics 51.5, 50.0.

Interspace hctween : First and second dorsals 18.3, 18.7 ; 2nd dorsal and
caudal 12.7, 13.9; base of pelvics and caudal 18.9, 20.4.
20 18

Teeth : — , —
27, 31

Description of type specimen. Trunk noticeably slender, its
height at pectorals (where highest) about 12 per cent as great as
its length to upper origin of caudal fin, its greatest thickness
about equal to its greatest height ; body narrowing rearward
with caudal peduncle nearly as thick as high. Head flattened
above, its length to origin of pectorals occupying about 33 per
cent of trunk to origin of caudal fin.

Snout obtusely wedge-shaped anteriorly, its length in front
of mouth about 50 per cent as great as length of head to origin
of pectorals, its length in front of eyes about 28 per cent. Eye
about 1.4 times as long as high, its horizontal diameter about 40
per cent as long as snout to mouth. Nostril with outer end
close to edge of snout; anterior narial flap triangular, with
pointed tip, reaching across narial opening. Pores on lower
surface of snout so small that they are visible only on close
scrutiny. Mouth low-arched, occupying about 80 per cent of
breadth of head. Preoral clefts short, extending about 33 per
cent the distance from corners of mouth toward nostrils. Fur-
rows from corners of mouth reaching rearward half way toward
first gill openings.

First gill openings a little shorter than distance between nos-
trils ; fifth openings a little shorter than first to fourth ; an-
terior margins only slightly concave, but tips of filaments ex-
posed in first to third. Teeth smooth edged as in other mem-
bers of this genus ; uppers with only 3 cusps, the axial much the
largest. Lowers with cutting edge nearly parallel with genera]
trend of the jaw. Cusp of the median lower tooth, identified as
such by its base overlapping that of the next tooth on either
side, directed strongly outward, to the one side or the other, on
all specimens seen. Two or three rows functional in upper jaw
hut one row only in lower jaw.

52 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Dermal denticles low, eonical-thorn-like, erect or nearly so
and so sharp that the skin feels very rough. Denticles on lower
surface straight, or nearly so, in random arrangement, but those
on sides and back slightly curved rearward and arranged in
single series in regular longitudinal rows (PI. 2) ; the rows on
the back anterior to the first dorsal fin converging rearward, but
the rows on the sides paralleling in general the main axis of the
trunk as far rearward as abreast the second dorsal, whence those
above the mid-level trend downward-rearward along the tail
sector and out along the caudal axis. The lower surface of the
trunk, like sides and back, is closely set with denticles, except for
the chin, the lips, a small area in the midline of the snout close in
front of the mouth, and the margins of the nostrils, which are
naked. Fins as a whole naked.

Base of first dorsal fin about as long as eye, its origin pos-
terior to the origin of the pectorals by a distance about as long
as from tip of snout to mouth. Rear end of base of first dorsal
about equidistant between perpendiculars at axil of pectorals
and at origin of pelvics. Distance from point of emergence from
skin of first dorsal spine to point of emergence of second dorsal
spine about as long as head to origin of pectorals. Second dorsal
spine about 1.8 times as long as first, each measured from point
of emergence from the skin, its tip reaching about 75 per cent
toward the upper corner of the fin. Distance from rear end of
base of second dorsal fin to origin of upper side of caudal fin
about 54 per cent as long as between points of emergence from
skin of first and second spines, and about 47 per cent as long as
length of head to pectorals; upper side of caudal fin about as
long as head to origin of pectorals, its shape as pictured in Fig-
ure 5A. Lower anterior margin of caudal 43 per cent as long as
upper margin. Distance from origin of lower side of caudal fin to
rear end of bases of pelvics about 75 per cent as long as head to
origin of pectorals. Rear end of bases of pelvics a little anterior
to point of emergence, from skin, of second dorsal spine. Pec-
torals truncate, with rounded corners, the distal margin, when
laid back, falling about abreast of point of emergence from skin
of first dorsal spine.

Tn their present state the edges of the pectorals, dorsals, and
pelvics, and the terminal sector of the caudal, are much frayed
out on all the specimens. But close examination makes it evident

BIGELOW AND SCHROEDER : SQUALOID SHARKS 53

that, normally, tliey are entire, not fringed as they are on *E.
schultzi (Bigelow, Schroeder, and Springer, 1953, p. 254, Fig.
9D).

Claspers of mature males not yet seen.

Color. Upper parts in general verj^ dark sooty gray, the mid-
line of the back paler (as a definite band on one of the speci-
mens), with a vaguely outlined pale yellowish spot on the top of
the head betw^een the eyes. Lower surface of head and body
sectors black; the black extending rearward in a narrow band
along the lower surface of the tail sector, nearly or quite as far
as the origin of the caudal ; also upward on each side close be-
hind the pelvics, to continue forward as a flank mark of the
shape shown on Figure 5A. The pattern of coloration, however,
is evident only on close examination, the sides, as a whole, being
very nearly as dark as the belly. Distal parts of the dorsal, pec-
toral and pelvic fins, and lower edge of caudal pale gray after
preservation, perhaps whitish in life.

The (presumablj^) luminescent system of markings to be seen
on various other members of Etmopterus is evident here only as
a row of black dots along the midline of the back from nape to
base of caudal. If similar markings are present elsewhere they
are masked by the dark hue and dense pigmentation of the sides.

Size. The size at maturity is not yet known.

Geographic range. So far known only off the northeast coast
of Florida, at the localities listed above (p. 50).

Remarks. The linear arrangement of the denticles on its back
and sides marks *huUisi off from all the other species of Etmop-
terus that were known previously from the North Atlantic and
Gulf of Mexico. It differs further from *spinax in its stouter,
thorn-like denticles, and in that the interspace between its first
and second dorsal fins is only about as long as from tip of snout
to first gill openings (as long as from tip of snout to origin of
pectorals in *spinax) ; from *hillianus and from *virens in its
noticeably shorter caudal peduncle, relative to the length of
the head; from *pusillus in its thorn-shaped denticles (truncate
with concave crown in ^pusillus) ; from *schultzi both in the na-
ture of the margins of its fins (p. 52) and in a relatively shorter
caudal fin (upper margin only as long as from tip of snout to
origin of pectorals in *bullisi, but as long as from snout to rear
edge of pectorals in * schultzi) ; from *princeps in a noticeably

54 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

longer interspace between its second dorsal and the origin of the
upper side of its caudal (to mention only the most conspicuous
difference) ; and from *polU in its tricuspid upper teeth (mostly
with 5 cusps in *polU), in the roughness of the lower side of its
snout, and in the minuteness of the pores there.

Among species known from other seas, it agrees with *lucAfer
(originally described from Japan) in the linear arrangement of
the denticles on the sides of the head and body, and in their
thorn-like nature, also with *villosus of Hawaiian waters; with
^hrachyurns, originally described from the Philippines (p. 59) ;
and with *aher7iathyi, from New Zealand. But there is no dan-
ger of confusing it with either one of these for the following
reasons: the distance from the tip of its snout to its first dorsal
fin spine is shorter relatively than in *villosus but its caudal fin
is considerably longer; the upper margin of its caudal is much
longer relatively than in *brachyurus ; the interspace between
its two dorsal fins (not longer than from tip of snout to first gill
openings) is noticeably shorter than in either *lucifer or *aher-
7iathyi (at least as long as from tip of snout to origin of pec-
torals), and its flank marks are much less conspicuous (cf. Fig.
5 A with Fig. 6A) than in either of these last two.

The Mediterranean shark, also, that was named Squalus in-
fernus by Blainville (1825, p. 59), a century and a quarter ago,
is referable to Etmopterus by Blainville 's account of its upper
teeth as three-cusped, but of its lowers as one-cusped, with hori-
zontal cutting edge." And there is nothing in the dimensions
given for it by Blainville to set it apart from *E. spinax, as ap-
pears from the accompanying comparative table. Our earlier
statement (Bigelow, Schroeder and Springer 1953, p. 224) that
its proportional dimensions differ from those of other Atlantic
species was based on a misreading of the measurements given for
it by Blainville.

Proportional dimensions in per cent of total length (A)
for the type specimen, 255 mm. ("dix pouces") long, of
inf emits Blainville 1825; and (B) for a female * spinax,
about 285 mm. long, from the Mediterranean, in the Mu-
seum of Comparative Zoology.

10 Notwithstanding its tooth characters, Garman (1913, p. 197) classed it as a
synonym of *CLntrophorus uyato Raflnesque 1810. We may also point out that
Blainville's (1825, p. 59) reference for it to his "PI. 14, Fig. 2" was incorrect;
iictually this figure represents a "Lmnna nnsua.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 55

A R

Snout to 5th gill openings 19 21

Snout to cloaca 57 58

Snout to 1st darsal fin 31 32

Snout to 2nd dorsal fin C3 63

Cloaca to lower origin of caudal fin 23 19

*S. spinax has been reported from southern Africa as well as
from the North Atlantic and Mediterranean, on the evidence of
a shark that was taken off Cape Point by the "Pickle" (Gil-
christ 1922, pp. 42, 49 ; Barnard 1925, p. 51 ; Smith 1949, p. 59),
with two others taken in the same general region by the "Dis-
covery" (Norman 1935, p. 37). And the "Pickle" specimen
certainly falls with *spinax in the relative positions of its fins,
as well as in the bristle-like nature and irregular distribution of
its dermal denticles, to judge from what little information is
available. But we still await a detailed description of it (or of
the specimens taken by "Discovery") ; nor are the outlines of
the black belly area and of the flank-markings shown on Smith's
illustration. Final decision, in short, as to the specific relation-
ship that the spinax-like form of South African waters bears to
the typical *spinax of the northeastern Atlantic must await a
comparison of specimens from the two ocean areas. None of the
other Atlantic species has been reported from farther southward
in the southern hemisphere that latitude 6°08'S {S. polli Big-
elow, Schroeder and Springer 1953, p. 241).

Other named sharks that can be definitely placed in Etmop-
terus are the following.

E. granulosus (Giinther) 1880, originally reported from the
southwest coast of South America, and subsequently from Ar-
gentina (Lahille 1921, p. 16, by name only) as well as from
southern Africa (Gilchrist 1922, p. 41; Barnard 1925, p. 49;
Smith 1949, p. 58). Barnard's reference of it to the Hawaiian
Islands, without supporting date, may actually have referred to
*E. villosus. The chief diagnostic features of granulosus are:
distance from tip of snout to emergence from skin of first dorsal
spine is at least as long as from emergence of first dorsal spine
to rear end of base of second dorsal (Giinther 1880, PL 2, fig.
C; Barnard 1927, PI. 2, fig. 8; Smith 1949, p. 58, fig. 50) ; dis-
tance from rear end of base of pelvics to origin of lower side of
caudal is shorter than from tip of snout to first gill openings;

56 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

margin of pectorals (when laid back) reaches nearly to abreast
of base of first dorsal spine; dermal denticles on sides of body-
described as granular, without median spine and in irregular
pattern, but those on tail in the form of minute spines and ar-
ranged in longitudinal bands (Giinther 1880, p. 19) ; body uni-
formly dark brown to blackish, but the fins edged with white
(Giinther 1880, p. 19), a feature not mentioned either by
Barnard or by Smith. Incidentally, the upper side of the caudal
fin is shown as about as long as from the tip of the snout to the
origin of the pectoral fins both by Giinther (1880, PI. 2, fig. C)
and by Barnard (1927, PI. 2, fig. 8), but only as long as from
snout to first gill openings by Smith (1949, p. 58, fig. 50), a
disagreement we cannot resolve.

*E. lucifer Jordan and Snyder 1902, Japan (Fig. 6 A-C). The
chief diagnostic features, as appear both from the original ac-
count and from our own examination of 5 specimens (two in the
Museum of Comparative Zoology, three in the U. S. National
Museum ) are : distance from tip of snout to point of emergence
from skin of first dorsal spine is about as long as from emergence
of first dorsal spine to midpoint of base of second dorsal ; upper
side of caudal and distance from rear end of base of pelvics to
origin of lower side of caudal both are about as long as from
tip of snout to origin of pectorals; rear end of base of first dor-
sal is about midway between perpendiculars at origin of pelvics
and at axil of pectorals ; dermal denticles slender, thorn-like,
weakly curved, sloping only slightly, and close set; those on the
sides arranged in general in longitudinal rows and directed
downward-rearward (a conspicuous feature), but those on the
black areas of the lower surface in random pattern, di-
rected obliquely downward-rearward on either side of the body
toward the midline, where a single row is directed rearward.
Upper surface of head, anterior to spiracles, with extensive
naked areas. Back and upper part of sides chocolate brown;
lower surface black, with a distinctive black marking on either
flank (Fig. 6A; Bigelow, Schroeder and Springer 1953, fig. 6B),
the line of transition from black to brown marked by a pale band
as shown on Figure 6 A. Imposed on the sides, there is also a
complex pattern of black dash-like markings, presumably
luminescent (Fig. 6B ; Oshima 1911, pp. 1-3), in longitudinal
linear arrangement.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 57

Originally described from Japan, Hucifer has since been
found in Sonth African waters (Gilchrist 1922, p. 49; Barnard
1925, p. 50; Norman 1935, p. 37; Smith 1949, p. 59, fig. 52).
It has also been reported by name from the Philippines (Smith
and Radcliffe 1912, p. 679, 21 specimens; Fowler 1941, p. 248).
and from Flores Straits in the East Indies (Weber 1913, p.
597). But we think it more likely on geographic grounds that
these reports from equatorial waters actually were based on *E.
hrachyunis (p. 58).

*E. villosus Gilbert 1905, from the Hawaiian Islands. The
combination of slender, spine-like denticles (those on back and
sides in linear series) with a first dorsal fin standing so far rear-
ward that its point of origin is about midway between the tip
of the snout and the origin of the upper side of the caudal fin
(corroborated by our examination of the type specimen in the
U. S. National Museum), sets ^villosus apart from all other
members of the genus yet known. Other features diagnostic of
*villosus are : upper side of caudal about as long as from tip of
snout to first-second gill openings; distance from rear end of
pelvics to origin of lower side of caudal fin only a very little
longer than from snout to level of spiracles; rear end of base of
first dorsal about midway between origin of pelvics and rear
margin of pectorals when the latter are laid back ; abdomen and
lower side of head blue-black, the sides brown, no black flank
marks being suggested by Gilbert's (1905, PI. 66) illustration
or by our recent examination of the type specimen; dorsals,
pectorals and pelvic fins broadly edged rearward with white ;
lower surface in general rough with denticles, the skin being
naked only on the lips and nostrils, close around the eyes and
spiracles, and close behind the dorsal, pectoral and pelvic fins,
which are partially so; the denticles along the back and on the
tail arranged in lengthwise series. On the type specimen, the
denticles along the midline of the lower surface are smaller than
they are pictured ; the first dorsal spine is only about 65 per cent
as long as the second ; the upper teeth have only three cusps ;
and the cusps of the low^er teeth are so strongly oblique that their
inner margins are nearly parallel with the jaw.

*Yillosus is so far known only from the type specimen, 170 mm.
long, taken by the "Albatross" off the south coast of Molokai
(Gilbert 1905, p. 580, PI. 66).

58 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

E. frontiniaculatus Pietschmann 1907, from Japan. In gen-
eral, this species resembles *E. lucifer in color, with brown back
and sides contrasting with ink-black belly and black flank marks.
But the latter differ in shape from those of Hucifer, as Oshima
(1911, p. 2, fig. 1) has pointed out, while neither Pietschmann 's
account nor his illustration (1908, p. 657, PI. 1, fig. 2) suggest
the presence of the linear pattern of luminous organs to be seen
on the sides of the body in Hucifer (p. 56). E. frontimaculatus
is set apart still more sharply from *lucifer by its denticles,
which (as pictured by Pietschmann 1908, p. 657, figs. 1, 2) are
truncate, with concave crowns (slender, thorn-like in *lucifer,
p. 56), while those on the sides are arranged irregularly (in
linear series in "^'lucifer). Also, according to Pietschmann (1908,
pp. 656-657) the skin of frontimaculatus is naked on the black
areas of the lower surface and of the tail, which in Hucifer
(p. 56) are as densely denticulate as the paler sides are and
the back. E. frontimaculatus resembles *E. pusillus of the At-
lantic in the nature of its denticles; Tanaka, in fact (1912, PI.
22, 1912^, p. 88) pictured and described a specimen from Misaki
under that name. But it differs sharply from *pusillus in the
nakedness of the black areas on its lower surface and on its tail ;
in a somewhat more rearward position of its first dorsal fin rela-
tive to the pectorals, and — more conspicuously — in its color
pattern, for the black belly and flank marks contrast with paler
brown sides and back which is not the case in * pusillus (com-
pare Pietschmann 1908, PI. 1, fig. 2 with Bigelow, Schroeder
and Springer 1955, p. 3, fig. 1). E. frontimaculatus is so far
known only from Japan.

*E. hrachyurvs Smith and Radcliffe 1912 from the Philip-
pines. This shark (Fig. 6 D, E) falls with Hucifer in the rela-
tive position of its pectoral, dorsal and pelvic fins ; in the thorn-
like nature of its dermal denticles (these roughen the lower sur-
face of its body as well as the upper) ; and in the arrangement in
longitudinal bauds of those on the upper surface, on the caudal
peduncle, and on the bases of the fins. It falls further with
Hucifer in its general color pattern, with sharply outlined black
flank markings, which are not shown on Smith and Radcliffe's
original illustration of it. But it differs from *lucifer in a rela-
tively much shorter caudal fin, the upper margin of which is
only about 66 per cent as long as from the tip of the snout to

BIQELOW AND SCHROEDER : SQUALOID SHARKS 59

the origin of the pectorals, as we can verify from recent examin-
ation of the type specimen in the U. S. National Museum (about
as long as from snout to pectorals in *lucifer). And there are
five cusps on its upper teeth (three in Hucifer), with the laterals
longer, relative to the median member, than in Hucifer. *E.
hrachyunis was originally described from an adult male, 227
mm. long, taken off Jolo Island in the Philippines by "Alba-
tross" in 1909, from 481 meters (263 fathoms). But its range
extends to southern Australia, provided our union with it of mol-
leri Whitley 1939 be correct (see below).

E. molleri (Whitley) 1939, from Australia. This shark de-
scribed by Whitley as Acanthidiuni molleri. seems not to differ
from E. brachyurus in any significant respect, to judge from
Whitley's description or illustration (Whitley 1939, p. 265, fig.
1; 1940, pp. 147-148, fig. 163).

E. haxteri Garrick 1957, from New Zealand. This newly de-
scribed Etmoptcrus, known from a female 742 mm. long, caught
at 914 meters (500 fathoms) seven miles south of Kaikoura,
N. Z., finds a close counterpart in E. princeps of the North At-
lantic. But it differs so sharply from princeps in the respects
summarized in the accompanying key (p. 63, alternatives 14-15),
that the two clearly represent distinct species.

E. ahernathyi Clarrick 1957, from New Zealand. This well de-
fined species, described from an immature male 338 mm. long
and from a female 278 mm. long, caught at 183-366 meters off'
Kaikoura, N. Z., falls in the subdivision of the genus that is
characterized by slender thorn-like denticles, with those on the
upper part of the sides arranged in unilinear, longitudinal rows,
and by a conspicuous pattern of black markings. Among the
other members of this division it comes the closest to brachyurus.
But it is separated from the latter by a much longer caudal fin
and by a more forward position of the first dorsal fin, to men-
tion only the most conspicuous differences (see Key, pp. 61, 62,
alternatives 7-10). For comparison Avith other members of its
geuus, we refer the reader to Garrick 's detailed account of aher-
nathyi and to his excellent illustrations of it.

The U. S. National Museum also contains a female Etmopterus,
about 270 mm. long (U.S.N.M. No. 91576) collected among the
Galapagos by the "'Albatross" in 1888, from 717 meters (392
fathoms), which agrees very closely with the West African

60 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

*polli, except that such of its upper teeth as are still intact have
only three cusps, contrasting with five cusps in *polli. Un-
fortunately the specimen is now in such poor condition that we
dare do no more than call attention to the presence in Gala-
pagan waters of a species of Etniopterus, as yet undescribed.

Finally, we face the case of E. paessleri Lonnberg (1907, p.
5, fig. 2) from the Straits of Magellan. Lonnberg 's reference of
this shark to Etniopterus has been accepted both by Regan
(1908, p. 43) and by Garman (1913, p. 229). And while Lonn-
berg states that lateral cusps were not visible ("nicht sichtbar")
on its upper teeth, the type (and only known) specimen was so
small (about 160 mm. long) that these structures may have been
overlooked. Until this point can be settled its generic status must
remain open. Meantime, we are content to let this species rest
provisionally in Etniopterus. Whatever the final outcome in this
regard, paessleri is set apart from all other members of Etniop-
terus by the nature of its dermal denticles, which are described
(they have not been pictured) as having a large central spine,
surrounded by several smaller spines ("einen grosseren Stachel
und mehere kleinere Nebenstacheln, " Lonnberg 1907, p. 5).
Also, its second dorsal fin-spine is described as a little smaller
than the first, just the reverse of the usual relationship.

Key to species of Etmopterus

1. Dermal denticles with larger central spine surrounded by several
smaller spines; second dorsal fin spine smaller than first dorsal spine;
lateral cusps not easily visible on upper teeth of small specimens

paessleri Lonnberg 1907.
Straits of Magellan, p. 60
Dermal denticles citiier truncate or with a central spine only, the
latter bristle-like or thorn-like ; second dorsal fin spine longer than
first dorsal spine; lateral cusps on upper teeth clearly visible, even
on very small specimens . . 2

2. Upper margin of caudal fin nearly as long as from tip of snout to
rear edge of pectorals when these are laid back *schult2i Bigelow,

Schroeder and Springer 1953.

Northern part of Gulf of Mexico, p. 50

Upper margin of caudal fin very little (if any) longer than from tip

of snout to origin of pectorals and considerably shorter in most 3

3. Dermal denticles on sides of body truncate, without central spine ... .4
Dermal denticles on sides of body with a central spine, either conical.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 61

thorn-like, or bristle-like 6

Back and upper part of sides dark brown, contrasting sharply with
a l)lack marking on either Hank, and with lower surface of head

and body; the black areas partially naked frontimaculattts

Pietschmann 1907. Japan, p. 58
Back and upper part of sides black or sooty like lower surface; no
conspicuous flank-markings ; lower surface, like upper, rough with

denticles 5

Denticles on sides of tail not arranged in regular longitudinal rows,
and of same shape as those on sides of body ; interspace between
first and second dorsal fins about as long as from tip of snout to axil

of pectorals *pusillus Lowe 1839. p. 49

Eastern Atlantic from equatorial
West Africa to the coast of Portu-
gal, including the Cape Verde
Islands, the Canaries, Azores and
Madeira; also Gulf of Mexico,
p. 49.
Dermal denticles on tail arranged in longitudinal rows, and more
spine-like than those on the body; interspace between first and second

dorsal fins shorter than from snout to first gill openings

granulosus Giinther 1880.
Coast of Chile, also reported from
Natal coast of southern Africa
and from Argentina, p. 55.

Denticles on sides in definite uniserial longitudinal rows 7

Denticles on sides in random arrangement, not in longitudinal rows . . 11
Upper side of caudal fin at least not longer than from tip of snout to

second gill openings 8

Upper side of caudal fin about as long as from tip of snout to origin

of pectorals 9

Distance from tip of snout to first dorsal spine is about as long as from
base of first dorsal spine to origin of upper side of caudal; interspace
between first and second dorsal fins is only about as long as from

tip of snout to level of spiracles *villosus Gilbert 1905.

Hawaiian Islands, p. 57.
Distance from tip of snout to first dorsal spine is little longer than from
first dorsal spine to rear end of base of second dorsal fin; interspace
between first and second dorsal fins is at least as long as from tip

of snout to second gill openings

*bradhyurus Smith and Eadcliffe
1912. Philippines; apparently in-
cluding E. molleri (Whitley) 1939,
southern Australia, p. 58.

62 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

9. Interspace between first and second dorsal fins is much shorter than
from snout to first gill openings ; black flank markings not con-
spicuous *hullisi, new species, off north-
east coast of Florida, p. 50
Interspace between first and second dorsal fins is as long as from tip
of snout to origin of pectorals, or longer; black flank markings
conspicuous .... 10

10. Distance from base of pelvics to origin of lower side of caudal is only
about as long as from tip of snout to first gill openings

*al)ernathyi Garrick 1957. New Zealand, p. 5
Distance from base of pelvics to origin of lower side of caudal is as

long as from tip of snout to origin of pectorals *lucifer

Jordan and Snyder 1902. Japan;
Atlantic off southern Africa; also
reported by name from Argentina,
the Philippines and the East In-
dies, p. 56.

For list of Philippine, East Indian,
and South African records, see
Herre 1953, p. 30.

11. Distance from rear end of base of pelvics to origin of lower side of
caudal is at least as long as from axils of pectorals to origin of

pelvics 12

Distance from rear end of base of pelvics to origin of lower side of
caudal is only about V2 as long as from axil of pectorals to origin of
pelvics 14

12. Distance from origin of pectorals to origin of pelvics is not longer
than from tip of snout to origin of pectorals *polli Bigelow,

Schroeder and Springer 1953. trop-
ical West Africa, p. 50
Distance from origin of pectorals to origin of pelvics is longer than
from tip of snout to origin of pectorals 13

13. Denticles on sides below first dorsal fin are slender, bristle-like (Bige
low, Schroeder 1948, Fig. 92B) ; skin on lower surface of snout and
in region of gill openings is rough with denticles; snout in front of
spiracles about as long as from spiracles to level of axil of pectorals ;
black flank markings as in Figure 5H

*}iillianus (Poey) 1861. Cuban and
West Indian region, and north-
ward along the American slope to
the offing of Chesapeake Bay.
p. 50.
Denticles on sides below first dorsal fin low, conical (Bigelow, Schroe-
der and Springer 1953, Fig. 10, D, E) ; skin on lower surface of snout

BIGELOW AND SCHROEDER : SQUALOID SHARKS 63

and in region of gill openings naked; snout in front of spiracles only
about as long as from spiracles to level of origin of pectorals ; black

flank markings as in figure 5G *viren.'i

Bigelow, Schroeder and Springer

1953. Northern part of Gulf of

Mexico, p. 50

14. Interspace between first and second dorsal fins is longer than from tip

of snout to axil of pectorals; distance from rear end of second dorsal

to origin of upper side of caudal is only about 35 per cent as long as

interspace between first and second dorsals

baxteri Garrick 1957. New Zealand, p. 59
Interspace between first and second dorsal fins is only about as long
as from tip of snout to origin of pectorals; distance from rear end
of second dorsal to origin of upper side of caudal is about 50 per cent

as long as interspace between first and second dorsals 15

1."). Dermal denticles on sides slender, bristle-like, and closely spaced (Fig.
3E) ; breadth of head about as great as length of snout to mouth;
longest gill openings only 14-33 per cent as long as eye; black flank
marks conspicuous, of shape shown in Figure 5F.. . .*spinax (Linnaeus)

1758. Eastern Atlantic, including
Mediterranean, from the Cape
Verde Islands, Morocco and the
Azores to Norway, p. 49 ; also
reported from southern Africa,
p. 56.
Dermal denticles on sides low, conical to thorn-like (more slender on
juveniles), and more loosely spaced (Fig. 3, F, G) ; breadth of head
1.2-1.4 times as great as length of snout (to mouth) ; longest gill
openings 33-50 per cent as long as eye; flank markings not conspicu-
ous, if visible at all *pnnceps Collett 1904. Faroes-
Hebrides region and offing of
Gibraltar in eastern Atlantic ; offing
of southern Nova Scotia to offing
of southern New England in west-
ern, p. 50.

Genus CexteOPHORUS Miiller and Henle 1837

Centrophorus Muller and Henle 1837, p. 398; 1937A, p. 115; 1841, p. 89;
type species *C. granulosus Miiller and Henle, Mediterranean, con-
sidered by them and by subsequent authors in general as the same as
Squalus granulosus Bloch and Schneider 1801, no locality given.

Generic Synonyms:

Squalus in part, Bonnaterre, 1788, p. 12, for *S. squnmosus Bonnaterre,

64 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

type locality not known; Bloch and Schneider 1801, p. 135 for *S.

granulosus Bloch and Schneider, no stated locality.
Acanihorlnnus in part, Blainville 1816, p. 121, for *A. squamosals Blain-

ville, equals Sqnalns squamosiis Bonnaterre 1788, and A. granulosus

Blainville, equals *Sqtialus granulosus Bloch and Schneider 1801.
Ccntrina in part, Cuvier 1817, p. 130, for C. squamosals Cuvier, equals

*Sqnalm squamosus Bonnaterre 1788.
Lcpidorhinus Bonaparte 1838, type and only included species, *Squ/ilus

squamosus Bonnaterre 1788.
Acanthias in part, Miiller and Henle 1841, p. 85, for A. uyatus Miiller and

Ilenle, equals *Sqiialus uyato Eafincsque 1810, Mediterranean.
Spinax in part, Bonaparte 1841, PI. 57, for Spinax uyatus Bonaparte, equals

^Squalus uyato Rafinesque 1810.
MaohephUus Johnson 1867, p. 713, type and only included species M.

duvi,eriU Johnson, Madeira.
Entoxychirus Gill 1862, p. 498, type and only included species E. uyatus

Gill, equals *Sqnalus uyato Rafinesque 1810.
Gaboa (subgenus of Centrophorus) Whitley 1940, p. 145, type and only

included species, *Centrophorus harrisonii McCuUoch 1915, Victoria,

Australia.
Som-nispinax (subgenus of Centrophorus) Whitley 1940, p. 146, type and

only included species, Centrophorus nilsoni Thompson 1930, New Zea-
land.
Probable Generic Synonym :
Atractophorus Gilchrist 1922, p. 48, type and only included species A.

armatus Gilchrist, off Natal, southern Africa. For discussion, see

p. 82.
Not Squalus Linnaeus 1758, type species S. acanthias Linnaeus.
Not Spinax Cuvier 1817, p. 129, type species Squalus acanthias Linnaeus

1758.
Not Acanthias Risso 1826, p. 131, type species A. vulgaris Risso, Mediter-
ranean, equals Squalus acanthias Linnaeus 1758.
Generic characters. Squalinae with the fin spines laterally
grooved and well exposed in most (but see p. 81 under C. nil-
soni), the second longer than the first; lower margin of caudal
with subterminal notch ; teeth one-cusped in both jaws, the
lowers wider than the uppers, with cusp directed strongly out-
ward all along each side of jaw, and with the bases of successive
teeth overlapping outwardly ; margins of upper teeth smooth ;
margins of low-er teeth finely serrate in some species but smooth
in others; the cusp of the median lower tooth either erect and
symmetrical or oblique ; upper teeth subtriangular, with the

BIQELOW AND SCHBOEDER : SQUALOID SHARKS

65

Fig. 8. Pectoral fin in diiferent species of Centropliorus, adjusted to ap-
proximately equal lengths along outer margin, to show shape of inner
posterior corner. A, foliaceus, same specimen as in Fig. 7F. B, squamosus,
female, 1230 mm. long, southwest of Iceland. C, stclndachneri, type speci-
men, after Pietschmann 1908, PI. 1, Fig. 2B. D, acws, type specimen, male
820 mm. long, Japan, Mus. Comp. Zool. No. 1049. E, uyato, male, 429
mm. long, northern part of Gulf of Mexico, Mus. Comp. Zool. No. 38165. F,
granulosus, female, 922 mm. long, northern part of Gulf of Mexico, U. S.
Nat. Mus. No. 157833. G, scalpratus, Victoria, Australia, after McCuUoch.

66 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

bases of successive teeth in contact along row, or slightly over-
lapping, the cusp erect and nearly symmetrical near center of
mouth in most species, but nearly or quite as oblique as the
lowers all along each side of jaw in a few species (p. 72) ;
median upper tooth either symmetrical or oblique ; inner corner
of pectoral fins at least quadrate and angular, and more or less
extended in most species (Fig. 8) ; snout to mouth in all known
species, not longer than from mouth to level of origin of pec-
toral fins ; nostrils without barbel ; preoral clefts present, more
or less expanded inwardlj^ (Fig. IB). Caudal peduncle without
preeaudal pits or lateral ridges ; dermal denticles on sides of
body low ; block-like, scale-like, or conical in different species
(Bigelow, Sehroeder and Springer 1953, p. 270, fig. 3).

Maximum recorded lengths, 1500 mm. for *C. granulosus and
1420 mm. for *C. squamosus (Bocage and Capello 1866, pp. 26,
27).

Depth range. The only definite records of depths of capture
that we have found are 631 meters for *foliaceus, off Japan
(Giinther 1887, p. 5), and 1756 meters in Philippine waters
(Smith and Radcliffe 1912, p. 679) ; 366-375 meters for * gran-
ulosus (specimen in U. S. Nat. Mus. from Gulf of Mexico) ; 50-
400 meters for *uyaio (Poll 1951, and specimens we have seen
from Gulf of Mexico) ; "common between about 400 and 1500
meters" for * squamosus (Grey 1956, p. 96) with one record
from 1875 meters; 400 meters for machiquensis (Maul 1955, p.
5 ; and 128 to 439 meters for scaipratus (Whitley 1940, p. 145).
While no definite information is available for any one of the
group of additional species described from Japan {^acus,
*atromarginatus, *tesselatus, and steindachneri) ,_ it seems that
their normal ranges are restricted to depths greater than 100-
200 meters.

Remarks. The most distinctive single character of the genus
(though probabl}^ not one of any great phylogenetic importance)
is the angular and usually more or less extended inner corner of
the pectoral fins. The range in this respect (Fig. 8) may be
illustrated by *C. foliaceus Giinther 1877 of Japan, in which the
pectoral inner corner is only veiy slightly extended or even
rectangular, through conditions in ^squamosus Bonnaterre 1788,
in *acus Garman 1906 (incl. steindachneri Pietschmann 1907),
in ^'uyato Rafinesque 1810, and in harrisonii McCulloch 1915,

BIGELOW AND SCHROEDER : SQUALOID SHARKS 67

where it is extended somewhat more, and in *granulosus Bloch
and Schneider 1801, where it is still further prolonged, as it also
is in scalpratus McCulloch 1915.

The gradation, in short, in the shape of the pectoral, is so
nearly unbroken that the revival of Lepidorhinus Bonaparte
1838 V Garman (1913, p. 211) and of Entoxychirus Gill 1862
by Fowler (1941, p. 242), based on differences in this regard,
serves no useful purpose, whether from the practical standpoint,
or as representative of probable phylogenetic relationship. In
the type specimens, in fact, of C. nilsoni from New Zealand
(Thompson 1930, p. 277), and of *C. tesselatus from Japan (now
in the Museum of Comparative Zoology; Fig. 10 C, D), the inner
corner of one of the pectoral fins is extended, but not that of the
other pectoral (apparently undamaged).

Teeth. The cutting edge of the lower teeth of *C. granulosus
is described and pictured both by Miiller and Henle (1841, p.
89, PI. 33) and by Bocage and Capello (1866, p. 26, PI. 1, fig.
3D) as finely serrate. Similarly, the reserve rows (5 in number)
of our eastern Atlantic specimen of this species are finely but
irregularly serrate in some places (Fig. 9A), though not in others.
The lowers in the more anterior of the two rows that are visible
from outside the mouth (no longer in function) also show faint
traces of serration here and there. But the lowers in the func-
tional row are slightly jagged in places, at most. And the situa-
tion is essentially similar in the case of a Gulf of Mexico
specimen of *granulosus, on which some of the low^ers show
microscopically fine serration, but others are merely somewhat
irregular in outline, probably as the result of wear. (For
further details, see Bigelow, Schroeder and Springer 1955, p. 6,
Fig. 2C, p. 8). Bocage and Capello (1866, PI. 1, Fig. 3B) pic-
ture the upper teeth of *granulosus as finely serrate, like
the lowers. But this seems to have been an error on the part of
the artist, for the uppers are smooth on both of the specimens
of *granulosus that we have examined — also on all the speci-
mens of other species of the genus that we have studied, for
that matter.

In *G. squamosus, as represented by a female about 1250 mm.
long from Iceland (Fig. lOA), and by jaws of a somewhat larger
individual, presumably from near the Faroes (Fig. IIB), the
lowers in the outermost of the two rows that are visible from

68

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Figure 9

BIGELOW AND SCHROEDER : SQUALOID SHARKS 69

outside the mouth are finely serrate along the central part of
the cutting edge, those in the reserve rows less clearly so. The
lowers in the first reserve row (the edges of the functional rows
are badly worn) are also unmistakably though faintly serrate in
places (Fig. 9C) on the type specimen of *tesselatus. On the
type specimen of *atromarginatus (on which the teeth are in
better condition), the lowers (Fig. 9D) are irregularly but un-
mistakably serrate. They are described as "serrated" in scalpra-
tus (McCulloch 1915, p. 98) from Australia also. And the two
rows of lower teeth that are visible from outside the mouth in
the type specimen of the Japanese *acus Garman 1906 show faint
and irregular traces of microscopically fine serration here and
there, though not so distinctly as these appear on Garman 's
(1913, PI. 12, fig. 6) illustration when the latter is viewed
under a lens. In the Australian harrisonii, also, the edges of the
lower teeth are irregularly serrate (Fig. 9E). On the other
hand, we have not detected any trace of marginal serration either
on the lower teeth of a juvenile male foliaceus, 351 mm. long,
from Japan (Fig. 7G), or on those of uyato, whether on a 429
mm. male from the Gulf of Mexico (Bigelow, Schroeder and
Springer 1953, p. 271, fig. 4) or on a 480 mm. male from the
Mediterranean. Thus the presence or absence of serrations on the
cutting edge of the lower teeth seems not to be a useful basis
for generic separation here, though it has long been so regarded
among the carcharhinid sharks.

Similarly, the degree of obliquity of the upper teeth ranges

Fig. 9. Lower teeth in different species of Centrophorus to show the
degree of marginal serration. A, graiiulosus, male, about 855 mm. long,
eastern Atlantic, Mus. Comp. Zool. No. 662, 2nd and 3rd reserve rows, x
about 9. B, sq^iamosus, third tooth, outermost row, from jaw of specimen
taken near the Faroes, Mus. Comp. Zool. No. 39571, x about 5.5. C, tess.e-
latus, male, 887 mm. long, type specimen, Mus. Comp. Zool. No. 1031, Japan,
first row visible from outside the mouth, x about 15. D, atromarginatus,
female about 870 mm. long, type specimen, Japan, Mus. Comp. Zool. No.
1455, first two rows visible from outside the mouth, x about 8. E,
harrisonii, Australia, teeth from one of McCulloch 's original specimens,
made available to us through the kindness of Dr. G. P. Whitley, x about
10. F, squamosus, lower teeth at center of jaw of same specimen as in B,
to show the obliquity of the median tooth, x about 2.4.

70

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

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71

Fig. 11. A, Ccntrophorus tessclatus, same specimen as in Fig. 9C, upper
and lower teeth, left hand side of mouth, x about 3. B, Centrophorus
squamosus, female, about 1100 mm. long, from southwest of the Faroes,
upper and lower teeth, left hand side of mouth, x about 1.3. C, Centro-
phorus squamosus, same specimen as in B, denticles from side below first
dorsal fin, x about 5.5. D, Centrophorns acus, male, 810 mm. long, type
specimen, Mus. Comp. Zool. No. 1049, Japan, denticles from side below first
dorsal fin, x about 15. E, Centrophorus atromargmatus, female, about 850
mm. long, type specimen, Mus. Comp. Zool. No. 1455, denticles from side
below first dorsal fin, x about 17. F, Centrophorus tesselatus, same speci-
men as in A, denticles from side, x about 17.

72 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

from the state in * granulosus and in *atromarginatus (Garman
1913, PI. 13, fig. 2), where those along the central sector of
the jaw are erect and symmetrical, or nearly so, to the state in
'*uyato (Bigelow, Schroeder and Springer 1953, p. 271, fig. 4)
and in harrisonii (McCulloch 1915, p. 99, PI. 14, fig. 3), where
this is true only of the first few teeth next to the midpoint of
the jaw, the teeth farther out along the latter being nearly or
quite as strongly oblique as the lower teeth are.

As remarked above (p. 21), the specimen on which Johnson
(1867) based his neAv genus and species MachepMlus dumerili,
but which Vaillant (1888, p. 70), Garman (1913, p. 212), Key
(1928, p. 440), and Bigelow, Schroeder and Springer (1953, p.
224) have considered as identical with *Centrophorus squamosus
(Bonnaterre) 1788, had a symmetrical tooth at the center of the
lower jaw. Accordingly Vaillant (1888, p. 70), followed by Rey
(1928, p. 443), included the presence or absence of a symmetrical
median lower tooth as alternative between two varieties of
*squamosus. Conditions in this respect among specimens of
Centrophorus that we have examined are as follows : *granulosus
(male about 855 mm. long, Europe, and female 922 mm.. Gulf
of Mexico) ; * squamosus (2 females about 1250 and 1280 mm.
long from Iceland and jaws of 4 somewhat larger specimens,
probably from west of the Faroes, Fig. IIB) ; *acus (type speci-
men, male about 820 mm., Japan) ; *atromarginatus (type speci-
men, 857 mm., Japan) ; and *foliaceus (male, 351 mm. long,
Japan), the cusp of the median lower tooth is about as oblique
as the cusps of the successive teeth outward, in each of the rows
that are visible from outside the mouth. On the other hand, the
cusp of the median lower tooth is erect and symmetrical in both
these rows on the type specimen of *C. tesselatus from Japan
(Fig. IIA). And the two specimens we have seen of *C. uyaio
are intermediate between these two extremes, the cusp of the
median lower tooth being strongly oblique on a male of 480 mm.
from Nice, France, but only very weakly so, as well as smaller
than that of the succeeding teeth on another male of 445 mm.
from the Gulf of Mexico (Bigelow, Schroeder and Springer
1953, p. 271, fig. 4). Evidently the degree of obliquity of the
cusp of the median lower tooth is not an appropriate basis for
generic separation here.

Dermal denticles. In some members of the genus the denticles

BIGELOW AND SCHROEDER : SQUALOID SHARKS 73

on the sides of the trunk are scale-like and overlapping, the rear
edges even or tridentate, and with the ridges on the outer surface
either parallel or diverging slightly rearward. In other species
the denticles are more block-like or conical-thorn-like, not over-
lapping, more or less pointed, with the ridges converging rear-
ward. And Key's (1928, p. 239, fig. 143) account of the denticles
of *C. granulosus as more pointed on juvenile specimens than on
adults seems actually to have been based on a specimen of *C.
uyato (Rey 1928, PI. 3, fig. 2), a species for which pointed
denticles are diagnostic. The shape of the denticles, indeed, is
the most obvious character among those likely to serve as alterna-
tive in this genus as between groups of species. But to subdivide
Ccntrophorus generieally on this basis, or by the other minor
characters on which Whitley (1940, pp. 145, 146) based his sub-
genera Gaboa (type Cenfrophorus harrisonii McCulloch 1915)
and Somnispinax (type Centrophorus nilsoni Thompson 1930),
seems not to serve any useful purpose, the total number of species
concerned being so small, and their resemblance, in general,
being so close one to another.

Nomenclatural status of the type species. The original descrip-
tion by Bloch and Schneider (1801, p. 135) of their Squalus
granulosus reads as follows: "Sq. bruneus cute granulosa, acu-
leis 2 robustis compressis, postice parum ineu ruis, intus con-
cavis, primo in initio pinnae dorsi, secundo in initio mesurae,
pinnis brevibus, dorsali pectoralibus propinquiorae quam ven-
tralibus remotissimis, mesura caudale vicina, foraminibus tem-
porum remotis, lunatis."

Obviously this is not enough to place Bloch and Schneider's
five-foot specimen, whether as to genus among the Squalinae, or
as to species, for it does not mention either the shape of the
pectoral fins or the nature of the teeth. It is therefore fortunate
for the sake of nomenclatural stability that while Miiller and
Henle's (1841, p. 89, PI. 33) description and illustrations of
their * granulosus were based on an alcoholic specimen from
near Sicily, they did have, for comparison, a dried specimen in
Berlin from Bloch 's collection, probably the one on which
Bloch and Schneider's 1801 account was based, for they mention
only the one. Miiller and Henle's identification of their
''granulosus" with the "granulosus" of Bloch and Schneider
1801 is therefore to be accepted — unless re-examination of the

74 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

type specimen (if still in existence) should eventually prove
otherwise. This, in turn, implies that Miiller and Henle's very
excellent description and illustrations take precedence over the
several accounts by subsequent authors, as the underlying basis
for the comparison of species within the genus Centrophorus.

Species. The following named species, listed in chronological
order, fall in Centrophorus as defined above (p. 64) : *Squalus
squamosus Bonnaterre 1788 eastern North Atlantic (Figs. IB;
SB; 9B; lOA, B; IIB, C), including Machephilus dumerili
Johnson 1867 ; *S. granulosus Bloch and Schneider 1801 includ-
ing Centrophorus hraganzae Regan 1906), type species, eastern
North Atlantic (including the Mediterranean) and Gulf of
Mexico; *8qualus uyato Rafinesque 1810, eastern Atlantic (in-
cluding the Mediterranean) and Gulf of Mexico (Fig. 8E) ;
Centrophorus moluccensis Bleeker 1860, Amboina ; Centrophorus
lusitanicus Bocage and Capello 1864, coast of Portugal ; *Centro-
phorus foliaceus Giinther 1877, Japan (Fig. 7F-I) ; *C. acus
Garman 1906, Japan (Fig. IID) ; *C. tesselatus Garman 1906,
Japan (Figs. 9C, IOC, llA) ; C. steindachneri Pietschmann
1907, Japan (Fig. 8C) ; *C atromarginatus Garman 1913, Japan
(Figs. 9D, HE) ; C. harrisonii McCulloch 1915, Australia (Fig.
9E) ; C. scalpratus McCulloch 1915, Australia (Fig. 8G) ; C.
nilsoni Thompson 1930, New Zealand ; and C. machiquensis Maul
1955, Madeira.

The specimen from southeastern Africa, described by Gil-
christ (1922, p. 48) as Atractophorus armatus, is also provision-
ally referred here to Centrophorus (for discussion, see p. 82).

The members of the genus may be segregated by the shape of
the denticles on the sides of the body, in three groups : A,
denticles block-like, close set but not overlapping; B, denticles
scale-like and overlapping more or less ; C, denticles spur-like or
conical-thorn-like and more widely spaced.

In the Atlantic, group A is represented by *C. granulosus,
by lusitanicus if this finally proves a distinct species (see below),
and by the newly described C. machiquensis, which maj^ prove
to be identical with '^granulosus when a larger number of speci-
mens of it have been examined; group B is represented by
* squamosus; group C by * uyato. The distinctive features of
*granidosus, ^squamosus and *uyato — all of which are fairly

BIGELOW AND SCHBOEDER : SQUALOID SHARKS

75

well known now — are discussed elsewhere (Bigelow, Schroeder
and Springer 1953, pp. 224-225; 1955, p. 7).

The status of lusitanicus remains uncertain. Originally de-
scribed as distinct from * granulosus by Bocage and Capello in
1864 but definitely classed by them in 1866 with *granulosus,
lusitanicus was revived as a separate species by Giinther (1870,
pp. 420-421),' and was so regarded both by Regan (1908) and
by Garman (1913, p. 199). Again relegated to the synonymy of
* granulosus by Rey (1928, p. 436), it has been revived by Nobre
(1935, p. 449). And Mr. G. E. Maul informs us that the first
dorsal fin of a specimen which he has examined in the British
Museum is larger than in ^granulosus, as appears from the
following tabulation.

Proportional dimensions, in per cent of total length, of (A)
Centropliorus lusitanicus, male, 742 mm. long, coast of Portugal,
based on data furnished to us by Mr. G. E. Maul from a specimen
in the British Museum, and (B) * Centropliorus granulosus,
female, 922 mm. long, Gulf of Mexico (U. S. Nat. Mus., No.
157833).

Snout length in front of movth
First dorsal fin : vertical height

Base, from anterior origin of spine
Free rear margin of fin
Second dorsal fin: vertical height

Base, from anterior origin of spine
Free rear margin of fin
Caudal fin: length of upper margin
Distance from snout to: origin of first dorsal spine
origin of second dorsal spine
upper origin of caudal fin
origin of pectoral fins
origin of pelvic fins
Interspace between : origin of first dorsal spine and
second dorsal spine

base of second dorsal and lower origin of
caudal
Distance from: origin of pectorals to origin of pelvics
origin of pelvics to lower origin of caudal

Until a larger number of specimens have been measured to
determine how wide the range of variation may be, it must

A

B

9.4

9.1

7.4

5.4

11.7

8.,5

7.3

6.9

6.1

5.0

6.6

5.7

4.0

4.5

19.7

20.3

32.1

34.8

67.0

69.2

80.3

79.7

22.2

22.2

60.7

58.7

34.9

34.4

6.6

6.0

38.4

37.6

15.8

18.1

76 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

remain an open question whether lusitanicus deserves recogni-
tion as a species separate from granulosus.

The situation is not clear for the species of Centrophorus
that are known from the northern Pacific (all of them from
Japan). Among these, group A (with block-like denticles closely
spaced in quincunx arrangement) is represented by *tesselatus
and by '*atroniarginatus. The demarcation between these two
seems clear cut, from our examination of the type specimens in
the Museum of Comparative Zoology. Thus the interspace be-
tween the rear end of the base of the first dorsal fin and the
point of emergence from the skin of the second dorsal spine is as
long as from tip of snout to axils of pectorals in *atromarginatus,
but is only as long as from the snout to origin of pectorals in
*tesselatus. Also, the inner corner of the pectoral fins is ex-
tended somewhat farther, the second dorsal spine is noticeably
longer, and the cusps of the lower teeth are not only more
strongly oblique but their cutting edge less deeply concave in
* atromarginatus than in *tesselatus. Furthermore, the cusp of
the median lower tooth, which is strongly oblique on the type
specimen of * airomarginatus, is erect and symmetrical on the
type specimen of *tesselatus, as noted above (p. 72). But this
last difference may be a sexual one.

It remains an open question whether *afromarginatus is sep-
arable from * granulosus of the Atlantic. At least, our compari-
son of the type specimen of the former (a female of 850 mm.)
with a male ^granulosus of 860 mm. from the eastern Atlantic,
with a female of that species, 922 mm. long from the Gulf of
Mexico (Bigelow, Schroeder and Springer 1955, fig. 2), and
with Miiller and Henle's (1841, PI. 33) excellent picture of a
^granulosus from the Mediterranean, has not revealed any dif-
ferences that might seem of specific rank, unless it be that the
gill openings are not as long in the Japanese form (third gill
is about 64 per cent as long as distance between nostrils) as in
our Gulf of Mexico specimen (third gill about 80 per cent as
long as distance between nostrils). And while Garman (1913,
p. 197) desscribed the teeth of * airomarginatus as "not ser-
rate," the lowers of the type specimen do show traces of fine
serrations along the cutting edge (Fig. 9D) much as in the case of
*granulosus (Fig. 9A), in *tesselatus also, for that matter (Fig.
9C). Pietschmann (1908, p. 663), in fact, has reported under

BIGELOW AND SCHROEDER : SQUALOID SHARKS 77

the name granidosns two Japanese specimens of Cenfrophorus,
respectively 536 mm. and 908 mm. long, that he had compared
with two granulosus of corresponding sizes from the Mediter-
ranean. But the actual identity of his Japanese specimens calls
for verification (if they are still in existence) for their denticles
are described (Pietsehmann 1908, pp. 663-667) as differing in
shape from those of ^granulosus of the Atlantic. Pietsehmann
also united both *tesselatus and *acus Garman with *granulosus.
But as implied in the foregoing discussion, *tesselatus differs
from * atromarginatus. And the denticles on the sides of the
body of *acus (no figures of which had appeared when Pietseh-
mann wrote) differ widely from those of * granulosus, for they
are scale-like, with tridentate rear margins (Fig. IID).

The *granulosus-*tesselatus-* atromarginatus group of the
Northern Hemisphere has a close ally in the Southern Hemisphere
of the Pacific in scalpratus of Australian waters, except that
McCulloch's (1915, PI. 13, figs. 2-7) illustrations show the denti-
cles on its sides as more pointed than those of * granulosus of
the Atlantic and Gulf of Mexico (Bigelow, Schroeder and
Springer 1953, p. 270, fig. 3A; 1955, fig. 2D), or of the Japanese
representative of the latter (p. 76). Other differences sug-
gested by McCulloch's illustrations of scalpratus may be due
to dift'erent methods of measurement, etc. The embryo from
Amboina that was described by Bleeker in 1860 as C. moluccensis
appears also to fall in this same group, for Regan (1908, p. 51),
who saw the type specimen, located it in the subsection of the
genus with "posterior angle of pectoral fin considerably pro-
duced and acutely pointed." And the little else that is known of
its fins and bodily dimensions is consistent with this. But the
shape of its denticles is not known.

Group B, with scale-like denticles (p. 74), is represented in
the northwestern Pacific by *foliaceus, originally described from
Japan, and reported by name (with very brief comment) from
deep water (960 fathoms) among the Philippines (Smith and
Radcliffe 1912, p. 679) ; by *acus from Japan; and by steindach-
neri, also from Japan. Among these, steindachneri, with tri-
dentate flank denticles and with the inner corner of the pectorals
only slightly extended (Fig. 80; Pietsehmann 1908, p. 676, text
fig. 3; PI. 1, fig. 1) agrees so closely with the type specimen of
•C. acus as to make it reasonably certain that the two represent

78 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

but a single species, to be named aciis on the ground of priority.
^'Acus, in turn, is allied to *foliaceiis. But it differs from the
latter in the evident extension of the inner corner of its pectoral
fins, for these are not appreciably extended in *foliaceus, either
as they are represented by Giinther (1887, p. 5, PI. 2, fig. A), or
as they appear on two immature males of *foHaceus, respectively
325 and 350 mm. long, from Japan (U. S. Nat. Mus., Nos. 161517
and 161518) that we have examined (Fig. 7F). The dermal
denticles, also, on the sides of the type specimen of *acus differ
conspicuously from those of *foUa,ceus, for while they are tri-
dentate in both cases, they are not only broader, relatively, with
shorter marginal teeth and less regularly overlapping in *acus
(Fig. IID) than in *foliaceus (Fig. 7H), but so much smaller
relatively that the one species is separable from the other at a
glance by this character alone. *C'. foUaceits, indeed, is by far
the roughest-skinned species of Centrophorns that we have seen.
*Acus is also firmer-bodied than *foliaccus, at least in the pre-
served state, and it is of a paler hue below than above, whereas
the specimens of *foliaceus we have seen are of as dark a choco-
late brown below as above. The species *acus may, therefore, be
retained as distinct, at least until someone re-examines the
type specimen of *foliaceus, now in the British Museum. (For
an account of *acus, with beautiful illustrations, see Garman
1913, p. 199, PI. 12, figs. 5-8.)

The accounts of *fol\aceus by Giinther (1887, p. 5) and by
Jordan and Fowler (1903, p. 631), do not especially emphasize
the characters now known to be diagnostic of species within the
genus Centrophorns. A more detailed account from this point
of view, with measurements, is therefore included here (p. 78).

Centrophorus foliaceus Giinther 1877
Figure 7F-I

Study material. Immature male, 351 mm. long, U. S. Nat. Mus.
No. 161518, Okinosa, Japan.

Proportional dimensions of specimen listed above, in per cent
of total length.

Snout, lengtli in front of: outer ends of nostrils 3.4; mouth 10.8.
Eye : length, 4.8.
Mouth: breadth, 7.6.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 79

Xostrils: distance between inner ends, 4.0.

GUI opening lengths: First, 2.1; second, 2.4; third, 2.4; fourth, 2.4; fifth

2.0.
First dorsal fin: vertical height, 5.0; length from imbedded base of spine

to rear end of base of fin, 9.0.
Second dorsal fin: vertical height, 5.0; length from imbedded base of spine

to rear end of base of fin, 8.0.
Caudal fin: upper margin, 21.1; lower anterior margin, 15.7.
Pectoral fin: outer margin, 10.8; inner margin, 7.7; width, 7.3.
Distance from snout to: imbedded base of first dorsal spine, 36.2; imbedded

base of second dorsal spine, 65.5; upper origin of caudal, 78.9; origin

of pectorals, 25.4.
Interspace between: rear end of base of first dorsal and imbedded base of

second dorsal spine, 22.5; rear end of base of second dorsal and

upper origin of caudal, 5.7; base of pelvics and lower origin of

caudal, 12.4.
Distance from origin to origin of: pectorals and pelvics, 33.7; pelvics and

caudal, 17.6.

16-1-16
Teeth :

14-14

Head, to orig-in of pectorals, about % of trunk to upper
origin of caudal fin; snout in front of mouth about 42 per cent
of head to pectorals; eye about 19 per cent as long as head;
distance from tip of snout to front of eye about 29 per cent of
head; distance between inner ends of nostrils about 37 per cent
as long as snout to mouth; longest gill openings (2nd, 3rd and
4th) about half as long as eye; mouth transverse, scarcely
arched; distance between inner ends of preoral clefts about 1.75
times as great as between nostrils.

Upper teeth short, triangular, the median tooth erect, sym-
metrical, but the subsequent teeth increasingly oblique outward
along the jaw, with the inner edges of those toward the corners
of the mouth nearly horizontal; the lowers wider than the up-
pers, the cusps directed so sharply outward as to form a nearly
unbroken cutting edge paralleling either side of the jaw. Cut-
ting edge of lowers perfectly smooth, both in the reserve rows
and in the outermost of the two rows visible from outside the
mouth ; the 2nd row slightly irregular, but not definitely serrate.
The condition of the specimen does not allow us to determine
whether or not there was a symmetrical median tooth in the
lower jaw.

80 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Dermal denticles scale-like, close set and largely overlapping,
those on upper and lower sides of snout ovate, but those on body
as a whole tridentate, lanceolate in general form, with the
median tooth much the longest, on short pedicle, rising steeply
from the skin and with high median crest; those on back, an-
terior to first dorsal spine, the largest. Length of first dorsal
fin, measured from anterior side of imbedded origin of spine
about 2.5 times as long as distance between nostrils; interspace
between rear end of base of first dorsal fin and anterior edge
of imbedded base of second dorsal spine about as long as from
tip of snout to second gill openings. Second dorsal fin similar
in shape to first dorsal, about 90 per cent as long, basally, as
first, similarly measured (see above). Dorsal spines both well
exposed; exposed portion of second about 1.5 times as long as
that of first ; distance between first and second spines, at points
of emergence from the skin, about 1.5 times as long as from
tip of snout to first gill openings. Interspace between rear end
of base of second dorsal fin and origin of upper side of caudal
about as long as from point of emergence from skin of second
dorsal spine to rear end of base of second dorsal fin. Caudal fin
about 20 per cent of total length, of the shape shown in Figure
7F. Distance from origin of lower side of caudal to rear end
of bases of pelvics about as long as from rear edge of eye to
origin of pectoral fins. Rear tips of pelvics (when laid back)
about in a line with point of emergence from skin of second
dorsal spine ; origin of pelvics anterior to imbedded base of
second dorsal spine by a distance about as long as from rear
end of eye to first gill openings. Pectorals with outer margin
about as long as from eye to fourth gill openings ; outer corner
broadly rounded ; distal margin weakly sinuous, reaching, when
laid back, about even with imbedded base of first dorsal spine ;
inner corner angular, with a few of the horny rays slightly ex-
tended (Fig. 8A).

Color. Plain dark brownish gray above and below; the rear
margins of pectorals and tip of dorsals pale edged, after many
years in alcohol.

Remarks. *C. foliaceus is recognizable at a glance, so widely
does its dermal armature, with close-set, strongly tridentate,
high-crested denticles, rising steeply from the skin, differ from
that of every other Centrophorus that we have seen, or that has

BIGELOW AND SCHROEDER : SQUALOID SHARKS 81

been pictured. Other distinctive characters for it (^Yithin the
genus Ceniropkorus) are: the shape of the inner corner of its
pectoral fins (p. 66), the lack of any abrupt transition from
the general contour of its back to the rising slope of the first
dorsal fin, and the pronounced obliquity of its teeth, uppers as
well as lowers, as described above (p. 79).

In all these respects the specimen here described agrees with
Giinther's (1887, p. 5, PL 2, fig. A) account of a 470 mm. (ISVo
inch) male, except that its inner pectoral corner is very slightly
extended ("rectangular, not produced" according to Giinther),
and that it is only the median upper tooth that is erect in the
upper jaw, the lateral upper teeth being strongly oblique (see
above), whereas Giinther characterized "the anterior teeth of
the upper jaw" as erect, the laterals as "slightly oblique."
These slight differences do not seem to call for recognition in
nomenclature on the basis of so few specimens.

The Japanese *foliaceus resembles *squamosus of the eastern
North Atlantic in its bodily proportions in general, in the rela-
tive locations of its fins, and in the scale-like nature of the
denticles on the sides of its trunk. But it differs from *squamo-
sus in the quadrate outline of the inner corner of its pectoral fins
(definitely though only slightly extended in *squamosus cf. Fig.
8A with Fig. 8B), and in the narrower, more regularly tri-
dentate and more steeply elevated denticles (cf. Fig. 7H with
Fig. IIC), so that the two species clearly are separable, the one
from the other.

The Northern Hemisphere species with scale-like denticles
are represented in the Southern Hemisphere by nilsoni of New
Zealand (Thompson 1930, p. 277, PI. 44). Nilsoni falls close to
*acus in the shapes of the inner corner of its pectoral fins and of
the denticles on the sides of its trunk. But its fin-spines are
described by Thompson (1930, p. 277) as "low and largely
covered," a character which (if normal) sets it apart.

The North Atlantic *uyato, with conical-thorn-like denticles
on its sides and with the inner pectoral corner moderately ex-
tended, is similarly replaced in the South Pacific by harrisonii
of Australian waters. Going back a step further in shark his-
tory, the embryo reported from Port Jackson, Australia, many
years ago by Steiudachner (1867, p. 333) under the name
Acanthias uyato Rafinesque, may have been the first specimen

82 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

of harrisonii to come under scientific observation. On the other
hand, Whitley (1934, p. 199) has suggested that it may have
been a Squalus, a view supported by Steindachner's description
of it as marked witli large white spots on its back. However this
may be, the cutting edge of the lower teeth, which are smooth
in the Atlantic *uyaio (p. 69), are irregularly serrate (Fig.
9E) in Jiarrisonii (described as imperfectly so by McCulloch
1915, p. 99). And other differences between the Atlantic and
Indo-Pacific populations may come to light when adequate series
of specimens from the two ocean areas have been compared.

Finally, we face the case of Atractophorus armatus from the
Natal coast of southern Africa, the type specimen of which
seems to have been a typical Centrophorus in all respects,
except that the tip of its second dorsal fin spine was described
and pictured as "barbed, like an arrowhead" (Gilchrist 1922,
p. 48, PI. 7, fig. 3). Unfortunately, the type specimen of
armatus is no longer to be found. ^^ But the collection of the
Government Marine Survey of South Africa does contain a
juvenile shark, 330 mm. long, apparently an armatus, the sec-
ond dorsal fin spine of which (while not two-barbed) bears a
single, wing-like expansion on its rear side, close to its tip (Fig.
7J). And any conformation of the spine tip, other than tapering
to a point (if normal) makes armatus unique among known
sharks, whether modern or fossil. We may either accept this
feature as justifying the generic separation of Atractophorus
from its parent genus Centrophorus, or we may transfer armatus
to Centrophorus, as the genus in which it would fall on the basis
of ancestral relationship, were it not for the single character in
question. We choose this last alternative, partly because we see
no useful purpose to be served in acting otherwise; partly be-
cause this choice seems the more likely of the two to focus
attention on armatus when additional specimens come to hand ;
and partly because of the chance that the peculiar conformation
of the second dorsal spine tip on the two specimens that have
been examined may have represented an abnormality, or may
have been the result of accidental damage.

If armatus is, in fact, a Centrophorus, its closest relationship
is with harrisonii, with which — and with *uyato of the North

11 We are so informed by Prof. J. L. B. Smith, who has made a special search
for it in the collection of the Government Marine Survey of South Africn.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 83

Atlantic — it shares pectoral fins with extended inner corner
(but shorter from origin to rear tip than from tip of snout to
level of first gill openings), and denticles on sides of body that
are described as acuminate, with many sharp ridges (Gilchrist
1922, p. 49).

Key to species of Centrophorus

1. Denticles on sides of trunk scale-like, overlapping, the ridges on their
outer surface (if more than one) parallel or diverging slightly rear-
ward 2

Denticles on sides of trunk block-like or conieal-thorn-like, not over-
lapping; the ridges on their outer surface converging rearvsrard, if at
all 5

2. Denticles on sides of body not definitely tridentate (Fig. IIC)

*squamosus Bonnaterre 1788. Northeastern Atlantic, p. 74
Denticles on sides of body regularly tridentate (Figs. 7H, IID) ... .3

3. Inner corner of pectoral fms quadrate, not appreciably extended
(Fig. 8A) ; dermal denticles on sides of body as in Figure 7H

*foliaceus Giinther 1877. Japan ;
also reported by name from the
Philippines (Smith and Radcliffe
1912, p. 679), and from New Zea-
land (Richardson 1956, p. 7). p. 78
Inner corner of pectoral fins definitely extended, though not greatly
so (Fig. 8D) ; denticles on sides of body as in Figure IID) 4

4. Both of the dorsal fin spines prominent and well exposed

aciis Garman 1906, Japan, p. 77

Both of the dorsal fin spines low and largely covered .

nilsoni Thompson 1930,
New Zealand; p. 81

5. Pectoral fin, from origin to inner rear corner, about as long as from
tip of snout to level of fourth gill openings scalpratus

McCulloch 1915, Australia, p. 74
Pectoral fin from origin to inner rear corner shorter than from tip
of snout to level of first gill openings 6

6. Denticles on sides of trunk block-like, c^uadrate, closely spaced in
quincunx arrangement; upper teeth erect and symmetrical, or nearly

so, all along central part of jaw 7

Denticles on sides of trunk conical-thorn shaped, loosely spaced in
random arrangement; upper teeth strongly oblique, except for the first
few next to the mid-point of the jaw, which are erect 8

7. Distance from rear end of base of first dorsal fin to point of emergence

84 BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY

from the skin of second dorsal spine is as long as from tip of snout
to axil of pectorals ; exposed part of second dorsal fin spine is about

75 per cent as long as distance between nostrils

* granulosus Bloch and Schneider
1801 (including lusitanicus Bocage
and Capello 1864) ; Mediterranean
with neighboring eastern Atlantic,
Gulf of Mexico, p. 73 ; and re-
ported by name from tropical West
Africa (Cadenat 1950, p. 101);
also machiqucnsis Maul 1955, Ma-
deira (p. 74) ; and atromargina-
tus Garman 1913, Japan (see dis-
cussion, pp. 76-77).
Distance from rear end of base of first dorsal fin to point of emergence
from the skin of second dorsal fin spine is only about as long as from
tip of snout to origin of pectorals; exposed part of second dorsal spine

only about 40 per cent as long as distance between nostrils

*tcsselatus Garman 1906, Japan, p. 74

8. Cutting edges of lower teeth smooth *uyato Eafinesque 1810.

Mediterranean, eastern tropical At-
lantic along the African coast from
Senegambia (Rochebrune 1883-
1885, p. 24) to Lat. about 20° S
(Poll 1951, p. 65); also Gulf of
Mexico, p. 74
Cutting edges of lower teeth irregularly serrate (Fig. 9E) . Jiarrisonii

McCulloch 1915. Australia, p. 74
Position uncertain : moluccensis Bleeker 1860, Amboina ; also armatus
Gilchrist 1922, southeastern Africa (p. 82).

Genus CeNTROSCYMNUS BocaRe and Capello 1864

Centroscymnus Bocage and Capello 1864, p. 263; 1866, p. 29; type species

*C. coelolepis Bocage and Capello, off Portugal.
Generic Synonyms :
Centrophorus in part, Giinther 1870, p. 423, for C. coelolepis Giinther,

equals * Centroscymnus coelolepis Bocage and Capello 1864; Thompson

1930, p. 277, for C. tvaitei Thompson, New Zealand.
Centroselachus Garman 1913, p. 206, type species *Centrophorus crepidater

Bocage and Capello 1864, off Portugal.
Proscymnodon (subgenus) Fowler 1934, p. 239, type and only included

species Centrophorus pluiiketi Waite 1909, New Zealand.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 85

Scymnodon in part, Fowler 1941, p. 225, for Centrophorus plunlceti Waite

1909, New Zealand.
Not Scymnodon Bigelow, Sehroeder and Springer 1953, p. 230, for Scymno-
don melas Bigelow, Sehroeder and Springer 1953, p. 233, fig. 5, which is
a young stage of Centroscymnus coelolcpis (p. 100-).

Generic characters. In fjeneral as in Centrophorus (p. 64),
except with inner corner of pectoral fins broadly rounded (this
is the most conspicuous difference between the two genera) ; the
upper teeth more slender than the loAvers, lancet shaped, those
along central sector of mouth nearly as long as those midway out
along either side of upper jaw; the successive teeth in the
functional row not in contact basally, or hardly so ; lower teeth
smooth edged, the successive cusps directed so strongly outward
as to form a nearly continuous cutting edge ; median lower tooth
not symmetrical ; second dorsal fin spine, like first, either ex-
posed more or less at tip, or entirely enclosed in the skin ; dermal
denticles on sides of body low, scale-like, with tridentate margins
on young specimens in some species, and probably in all, also
on adults in some species, but replaced later in growth in other
species by denticles with evenly rounded margins and concave
crowns ; preoral clefts short in some species but so long in others
as to leave only a narrow gap between their inner ends (see dis-
cussion, p. 87), considerably expanded inwardly in species
where they are short (Fig. IC) but not appreciably so in those
where they are longest (Fig. ID).

Maximum recorded length 1140 cm. for C. coelolepis (Bocage
and Capello 1866, p. 31) and 1414 for C. plunketi (Waite 1910,
p. 384).

Depth range. Centroscymnus coelolepis (p. 88), the shoalest
record for which has been 329 meters, has its center of distribu-
tion between 400 meters and 2000 meters according to locality,
and has been taken down to about 2700 meters (maximum yet
recorded for anj- shark ) . It ma}- fairly be classed as the deepest
living of modern sharks. (For further details see Bigelow
and Sehroeder 1948, p. 498, and Grey 1956, pp. 97-98.) Depth
records for the other members of this genus are 270-920 meters
for crepidater off Iceland (Saemundsson 1922, pp. 192, 196,
as Centrophorus jonsonii) ; 220 meters for plunketi (Waite
1910, p. 384) ; 780 meters for rossi (Alcoek 1898, p. 144) ; and
512 meters for fiiscus (Gilchrist and von Bonde 1924, p. 3).

86 BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY

Remarks. Teeth. The only characters mentioned by Bocage
and Capello (1864, p. 263;' 1866, pp. 19-20, 24, 29) in their
successive generic diagnoses of Centroscymnus that contrast with
their diagnosis for Centrophorus (1866, pp. 19, 22) were that
the upper teeth of the former are awl-shaped and loosely spaced
("subuliformes ecartes") and that its first dorsal fin spine is
almost entirely enveloped by the skin. But a much more con-
spicuous difference than either of these is the rounded inner
corner of the pectorals of Centroscymnus, as contrasted with
their angular inner corner in Ceritrophorus (p. 66). Similarlj^
the onlj^ alternative character in their original diagnosis of
Centroscymnus, as contrasted with Scymnoclon, is the shape of
the lower teeth, which Bocage and Capello characterized for the
former as not unlike those of CentropJiorus (i.e. as oblique,
without symmetrical median tooth), but for Scymnodon as with
symmetrical median tooth, the first tooth outward from it on
either side erect, the succeeding teeth as increasingly oblique
toward corners of mouth ("dens medianus impar, post eum
dantes premum erecti, deinde versus angulum oris magis ac magis
decumbentes, " Bocage and Capello 1864, p. 263). But a second
difference between the type species of Centroscymnus (*coelo-
lepis Bocage and Capello 1864) and of Scymnodon {* ring ens
Bocage and Capello 1864), equally diagnostic and much more
conspicuous, is that while the upper teeth along the central
sector of the mouth are nearly as long as those midway out along
each side of the jaw in Centroscymnus, they are much shorter
than those midway along the jaw in Scymnodon. The value of
this feature as a diagnostic character seems first to liave been
realized by Key (1928, p. 486, fig. 152).

Dermal denticles. In the adults of some species {plunketi
Waite 1910, New Zealand; *crepidatcr Bocage and Capello 1864,
eastern Atlantic) that are referable to Centroscymnus by their
dentition, the denticles on the sides of the trunk are tridentate
in outline, the outer surface sculptured with longitudinal ridges.
In the adults of others {*coclolepis Bocage and Capello 1864,
both sides of the North Atlantic; cryptacanfhus Regan 1906,
Madeira; *owstoni Garman 1913, Japan), they are rounded in
outline, with smooth, concave crowns; also in fiiscus Gilchrist
and von Boude 1924, southern Africa, so far as the concavity

BIGELOW AND SCHROEDER : SQUALOID SHARKS 87

of the crowns is concerned/" But it has been found by Tortonese
(1952, p. 386, corroborated by Bigelow and Schroeder 1954, p.
47) that the rounded denticles of the adult *coelolepis, and ap-
parentW those of *owstoni as well (Bigelow and Schroeder 1954,
p. 50), are preceded at early stages in growth (as proven by
embryos of known parentage) by trideutate forms, the replace-
ment of denticles of the juvenile shape by others of the adult
shape taking place when a given specimen is half to two-thirds
grown. For further details we refer the reader to our earlier
account (Bigelow and Schroeder 1954, p. 47, fig. 2). It follows
from this that our earlier acceptance of the shape of the denticles
(Bigelow and Schroeder 1948, p. 451, following Garman 1913,
p. 189) as alternative between Centroscymnus, where they are
rounded on the type species at maturity {*coelolepis Bocage and
Capello 1864) and Scymnodon, where they are tridentate at
maturity, is no longer tenable.

It seems appropriate here to recall that we owe to Steen-
strup (1861) our knowledge that the dermal denticles of sharks
do not persist throughout the life of the fish, growing as the
latter grows, as the scales of bony fish do, but that they are
constantly being shed, to be replaced by other, larger denticles.
Thus several generations of denticles succeed one another dur-
ing the life-span of any individual shark.

Fill spines. It seems worth emphasizing that the imbedded
basal part of the fin spines is easily felt, in species in which
their tips are concealed by the skin (p. 89).

Preoral clefts. The preoral clefts extend for about 40-50 per
cent of the distance from the corner of the mouth toward the
midline of the snout in *coelolepis (incl. cryptacanthus) of the
North Atlantic, in '*owstoni of Japan, and in macracantJnis of
the Magellanic region ; about 60 per cent of that distance in
waitei from New Zealand;^" but for about 90 per cent of that
distance in *crepidater (incl. *jonsonii) of the North Atlantic
(Fig. ID) and in rossi Alcock 1898 of the Indian Ocean (Alcock
1899, PI. 26, fig. 3). Both *crepidater and rossi fall, however,
with the type species of Centroscymnus {* coelolepis) in dental
characters, in shape of pectoral fins, and in shortness of fin-

12 The contour of the margins of the denticles is not known for fuseus.

13 Taken from a drawing made available to us by Dr. J. A. F. Garrick before
publication.

88 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

spines. Hence the difference in the length of the clefts does not
call for generic recognition, especially where the total number
of species concerned is so small.

Shape of caudal fin. Conditions in Centroscymnus argue
against the presence or absence of a subterminal notch on the
loAver margin of the caudal as a subfamily alternative among
the Squalidae, for while this notch is well marked in most of
the members of the genus, it is indistinct in waitei (Garrick
1955, p. 233, fig. lA). This seems the appropriate place for
us to point out that the caudal of *coelolepis is not correctly
pictured in our earlier illustration (Bigelow and Schroeder
1948, fig. 94). Actually it is of the shape shown here on Figure
12 H.

Species. The following named species, listed in chronological
order, are referable to Centroscymnus as defined here : *Centro-
scynmus coelolepis Bocage and Capello 1864, type species, North
Atlantic, east and west; *Centrophorus crepidater (Bocage and
Capello 1864), off Portugal; Centrophorus rossi Alcock 1898,
Travancore Coast, India; Centroscymnus cryptacanthus Regan
1906, Madeira ; Centroscymnus macracayithus Regan 1906, Magel-
lanic region; * Centroscymnus owstoni Garman 1906, Japan;
CeyiiropJwrus plunketi (Waite) 1910, New Zealand (referred by
Garman 1913, p. 210 and by Fowler 1941, p. 228 to Scymnodon
because of its tridentate denticles) ; * Centrophorus jonsonii
(Saemundsson) 1922, Iceland; Centroscymnus fuscus Gilchrist
and von Bonde 1924, southern Africa ; Centrophorus waitei
(Thompson) 1930, New Zealand."

Detailed descriptions of the type species, *coelolepis, with
illustrations, have been given by Garman (1913, p. 204, PI. 17,
figs. 5-8), by Rey (1928, p. 451) and by Bigelow and Schroeder
(1948, p. 494; 1954, p. 47). Coelolepis is known in the eastern
Atlantic from the offing of Cape Verde and of Morocco to the
Faroe Bank and to Iceland, including the western part of the
Mediterranean ; from the offing of Delaware Bay to the Banks of
Newfoundland in the western. According to Regan's (1906, p.
437) account, his cryptacanthus from Madeira differs from the
specimens of * coelolepis that we have seen (including the one

M Our inclusion of lonitei in CentroscymnuB follows Garrick's (1955) study of
the type specimen, ttie result of which was contributed to us by Dr. Garricli
in advance of his publication.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 89

on which Garman's illustration was based) in a slightly longer
snout; in an interspace between the two dorsal fins about 6
times as long as the base of the first dorsal (4-5 times in coelo-
Ic-pis) ; in the extension rearward of the tips of the pelvic fins
to abreast of the tip of the second dorsal fin; and in complete
enclosure of the fin spines by the skin. Also, the denticles on
the trunk rearward from abreast the first dorsal fin are described
(Regan 1906, p. 437) as already rounded, and with concave
crowns on a cryptacanthus only 700 mm. long (the type speci-
men), whereas various transitional stages are represented on a
*coelolepis of about that same size (690 mm., Bigelow and
Schroeder 1954, p. 48, fig. 2). But with both *coelolepis and
cryptacanthus recorded from Madeira, the desirability is self-
evident of comparison of the type of cryptacanthus (in the
British Museum) with northern specimens of *coelolepis.

*C. owstoni from Japan (the type specimen of which is in the
Museum of Comparative Zoology) differs from *coelolepis of the
Atlantic in a considerably longer snout, a somewhat larger eye,
a considerably larger second dorsal fin relative to the size of
the first dorsal, with the rear tip reaching more nearly to the
upper origin of the caudal fin, and in relatively smaller —
consequently more numerous — dermal denticles on the sides
of the trunk.

The New Zealand shark that was described by Waite (1910)
as Centrophorus plunketi, but which has been referred by Gar-
man (1913 p. 210) to Scymnodon, and by Fowler (1933, p. 239;
1941, p. 228) to his subgenus Proscyinnodon, is set apart from
the coelolepis-owstoni pair hy the tridentate shape of the denti-
cles on the sides of the trunk on large specimens (type specimen
a female 1414 mm. long) — presumably on small specimens also.
And its denticles are both described and pictured (Waite 1910,
p. 386, text fig.) as having as many as eight separate roots. The
tips, also, of its pelvic fins are pictured (Waite 1909, PI. 37) as
reaching back only about as far as a perpendicular at the point
of emergence of the second dorsal fin spine (they reach to a
perpendicular at about the midpoint of the free lower margin
of the second dorsal fin both in *coelolepis and in * owstoni),
while the second dorsal fin is pictured as considerably larger in
plunketi than it is either in *coeloIepis or in * owstoni.

Dr. J. A. F. Garrick's study of the type specimen (a juvenile

90 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

male, 318 mm. long) of the shark described by Thompson (1930)
as Scymnodon waitei places it definitely in the genus Centro-
scymnus as here defined, while his detailed account of it, with
excellent illustrations, makes possible, for the first time, a satis-
factory vicAv of such of its characteristics as are the most
evidently diagnostic of the species. Briefly, these are : snout in
front of mouth shorter than distance from mouth to level of
first gill openings; head, to origin of pectorals, about 20 per
cent of total length ; distance between inner ends of preoral
clefts nearly or quite as long as between nostrils; base of
second dorsal about twice (1.9) as long as base of first dorsal;
dorsal fin spines with only the tips exposed; caudal fin without
subterminal notch ; dermal denticles on sides of body strongly
tridentate, on four-angled base, and rising steeply from the
skin ; the outer exposed surface concave, M'ithout axial ridge ; the
denticles on the lower side of the snout ovate, with pointed tip
and with prominent axial ridge. (For further details, including
teeth, see Garrick 1955, pp. 233-238, figs. 1, 2.)

Thompson (1930, p. 278) followed by Richardson and Garrick
(1953, p. 35) has suggested that waitei may prove to represent
a stage in the growth of Centroscymnus j)lunketi, intermediate
between the adult female that was the basis of Waite's original
(1910) account of the latter, and the embryo, 165 mm. long, that
was pictured subsequently by him (Waite 1914, p. 127, PI. 3),
from a litter of 36 taken from a female measuring 1398 mm.
Solution of this question is, however, a matter for the future.

On the Magellanic shark named C. macracanthus by Regan
(1906, p. 436), the dorsal fin spines (first as well as second, Fig.
12 E, F) project farther from the skin than they do in any of
the other representatives of the genus discussed so far, though
not as much as is suggested by Regan's (1906, p. 437) character-
ization of them as "strongly projecting." Its two dorsal fins
also — especially the first — are larger relatively than in the
specimens of *coelolepis and of *owsioni that we have seen, more
nearly paralleling plunketi in this respect.^^ And the distance
between the inner ends of its preoral clefts is nearly as long as

15 We are indebted to Dr. Ethelwyn Trewavas and to Mr. Denys W. Tucker of
the British Museum (Natural History) for tracings of the first and second dorsal
tins, and to Dr. Ethelwyn Trewavas for the accompanying drawinir of a lower
tooth (Fig. 12 G), and for other details of the type specimen.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 91

between the inner ends of the nostrils, to judge from Regan's
(1906, p. 436) statement "length of anterior labial fold about
equal to its distance from the symphysis." The flank denticles
are tridentate on the type (and only known) specimen, but
the latter is so small (640 mm. long, Regan 1906, p. 437) that
this may represent a transitory state ; those on mature specimens
may be of a different shape, as they are in *C. coelolepis (p.
87).

In the species of Centroscymnus so far mentioned, the gap
between the inner ends of the preoral clefts is at least % as long
as the distance between the nostrils. In the three named species
yet to be considered {*crepidater and *jonsonii of the northeast-
ern Atlantic; rossi of the Indian Ocean) the clefts are so much
longer that the space between their inner ends is only about one-
third as long as between the nostrils (Figs. ID, 12B). Other
than this, the most sharply diagnostic features of *crepidater
are that its flank-denticles are tridentate on large specimens
(Fig. 12D), as they also are in C. plmiketi (p. 89), and that
its first dorsal fin spine projects from the skin nearly as far as
in C. macracanthus (cf. Fig. 12A Avith 12E, F).

The original (and only detailed) account of *G. jonsonii
(Saemundsson 1922, p. 192, PI. 5) does not include any com-
parison with *crepidaier, nor does it suggest any definite ground
for considering it as distinct from the latter. Comparison of
seven specimens (male and female, 587-796 mm. long) from
south of Iceland and southwest of the Faroes ^"^ with the 787 mm.
female from Madeira (Fig. 12A), on which Giinther (1870, p.
421) based his first-hand account of *crepidater,^' has revealed
no significant differences, whether in teeth, in the shape of the
denticles on large specimens, in bodily proportions ( p. 93),
or in the shapes, the sizes or the relative locations of fins. Con-
sequently we have no hesitation in classing * jonsonii as a syn-
onym of *crepidater, the known range of which extends (on this
basis) from Madeira and the coast of Portugal northward to the
Faroes and southern Iceland.

16 Kindly made available to us by the Natural History Museum, Reykjavik,
and by the University Zoological Museum, Copenhagen.

1" We owe to the British Museum the opportunity to examine this specimen.
Its identification as *crepidater has been checlsed through correspondence with
Dr. A. M. Kamalho, who has contributed measurements and photographs of the
teeth of the type specimen which he has examined in Lisbon on our behalf.

92

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

r-l

O

BIGELOW AND SCHROEDER : SQUALOID SHARKS 93

Dr. J. A. F. Garrick informs us, also/^ that the Canterbury
Museum contains two specimens of a shark taken on a long line
set at 914 meters (500 fathoms) off Keikoura, New Zealand,
and two others presumably from New Zealand but without
definite locality, that differ from *crepidaier (including *jon-
sonii) of the eastern North Atlantic only in such minor partic-
ulars that it is doubtful Avhether specific separation is warranted.
And this we can corroborate, for we find no significant differ-
ences in bodily proportions, fins, teeth or denticles between one
of these New Zealand specimens (kindly sent us by Dr. Garrick)
and the Atlantic specimens that we have seen of *crepidater
(pp. 93-94).

Proportional dimensions in per cent of total length of *Cen-
troscymnus crepidater. A, female, 787 mm., Madeira, from the
British Museum (Natural History) ; B, four females, 746 to 796
mm. long and C, male, 587 mm. long, off Iceland and the Faroes,
from the University Zoological Museum, Copenhagen ; D, female,
822 mm. long, presumably from New Zealand (see above), Mus.
Comp. Zool. No. 39570.'

Fig. 1:2. A, Centroscymnus ci-epidater, female, 787 mm. long, from
Madeira, in British Museum. B, lower surface of head of same, to show
the great length of the preoral clefts. C, upper and lower teeth of same, x
about 2.3. D, denticles of same, from side below first dorsal fin, x
about 6.5. E, Centroscymnus macracanihiis, outline of first dorsal fin of a
female, 640 mm. long, type specimen, in British Museum, from Magellanic
region. F, outline of second dorsal fin of same; E and F, after tracings
of the. fins kindly contributed by Mr. Denys W. Tucker. G, third right
hand lower tooth of same, x about 7, from drawing kindly contributed by
Dr. Ethelwyn Trewavas. H, Centroscymnus coelolepis, caudal fin of
female, 1035 mm. long, from offing of Delaware Bay, Mus. Comp. Zool. No.
38295.

18 Information contributed by Dr. Garrick in advance of publication.

94

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

A

B

C

D

Trunk at origin of pectoral : breadth

8.9

8.3-9.3

8.7

9.0

height

7.4

6.7-8.8

8.2

7.1

Snotit in front of : eye

8.6

8.2-9.3

8,3

8.9

mouth

11.4

11.4-12.7

11.7

11.8

Eye: horizontal dianietpr

4.4

4.4-4.6

4.6

4.5

Mouth: breadth

6.1

6.1-7.5

7.5

7.1

Xostrilft: distaiiee between inner ends

3.3

3.1-3.6

3.2

3.6

Preoral clefts: distance between inner ends

1.1

1.1-1.3

1.0

0.7

(iill opening lengths: first

1.5

1.3-1.5

1.4

1.3

third

1.5

1.3-1.5

1.4

1.3

fifth

1.5

1.3-1.8

1.6

1.6

First dorsal fin: vertical height

5.2

5.1-5.5

5.4

4.6

length of base from origin of spine

7.5

5.9-6.7

7.0

6.6

Second dorsal fin: vertical height

4.8

4.3-5.9

5.4

5.0

length of base from origin of spine

7.1

6.3-6.8

7.0

7.3

Caudal fin : upper margin

oo o

19.0-20.7

20.8

18.9

lower anterior margin

13.2

10.7-13.5

13.0

13.4

Pectoral fin : width

5.3

5.2-6.8

4.2

5.2

length of anterior margin

13.1

11.8-13.1

11.4

12.4

Distance from snovt to : first dorsal

28.9

28.1-30.5

30.7

29.2

second dorsal

60.2

61.0 04.2

60.3

61.7

upper caudal

77.8

79.3-81.5

79.2

81.1

pectoral

24.4

23.7-24.1

24.7

23.3

pelvics

60.6

61.0-65.0

.16.3

62.2

Interspace between: origins of first

and second dorsal spines

30.9

28.0-31. .-;

32.0

29.5

second dorsal and caudal

G.l

6.9-8.0

7.0

7.7

pelvics and caudal

8.0

7.4-8.8

lO.O

7.9

Distance from origin to origin of :

pectorals and pelvics

36.1

36.6-40.7

31.7

40.5

pelvics and caudal

15.8

15.2-17.9

17.9

15.7

Teeth, number of series : uppers

43

39-48

51

36-f^'>

lowers

36

32 36

32

364-1='

This adds to the list of cases among sharks and skates where
a species, or pair of very closely allied species, is known from
temperate latitudes in the two hemispheres, hut not from the
tropical-subtropical belt that intervenes.

C. rossi from the Indian Ocean, if represented correctly in
Alcock's (1899, PI. 26, Fig. 3) illustrations, differs from *crep-
idater in a much longer head, and in detail as to the vinpaired
fins, especially the caudal which appears to lack the subterminal

19 Several teeth have been lost at the corners of the mouth.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 95

notch ; perhaps also in the shapes of the teeth, though the illus-
trations of the latter by Alcoek appear to be someAvhat diagram-
matic. The denticles on the sides of the trunk are stronglv
tridentate on juveniles (represented by the type specimen 254
ram. long), much as they are in *C. coelolepis (Bigelow and
Schroeder 1954, p. 48, Fig. 2B) ; the shape is not known for
adults.

The relationship of C. fnscus of southern African waters to
other members of its genus remains doubtful, for while it falls
with *coelolepis in length of snout, in the sizes and relative posi-
tions of fins, and in the concave crowns of the denticles on the
sides of its body when mature (type and only known specimen
1,100 mm. long),^° the length of its preoral clefts is not known.
And the characterization of it in the original description (Gil-
christ and von Bonde 1924, p. 2, our only source of information)
as with orbits 1.7 times as long as the snout (''orbit .6 in
snout") seems an obvious error, for this would credit it with
eyes far larger than those of any related shark.

Key to Species of Centroscyninus

1. Distaiue between inner ends oi preoral clefts at least 7.j per cent as

long as distance between nostrils 2

Distance between inner ends of preoral clefts is not more than abont
30 per cent as long as distance between nostrils C

-. Exposed part of lirst dorsal fin spine, measured along its anterior
margin, is very nearlj- 50 per cent as long as distance from its point of
emergence from tlie skin to rear end of base of the fin (Fifj. 1'2F) . .

macracanthus Began 1906.

Magellanic region, p. 90

Dorsal tin spines exposed only at their extreme tips, if at all ii

3. Both of the dorsal fin spines are wholly concealed by the skin

cryptacantlius Regan 1906.

Madeira, p. 88

The tips of the dorsal fin spines protrude more or less from the skin . .4

4. First dorsal fin, from point of emergence of spine to rear end of base,
is about as long as second dorsal fin, similarly measured . ^coelolepis

Bocage and Capello 1864.

North Atlantic, east and west. p. 88

First dorsal fin, from point of emergence of spine to rear end of base.

is shorter than second dorsal fin, similarly measured r>

20 .Apparently the tj'pe specimen has been lost (Smith 1949, p. 58).

96 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

5. Snout in front of mouth is at least as long as from mouth to level of
first gill openings *owstoni Garman 1906.

Japan, p. 89
Snout in front of mouth is noticeably shorter than from mouth to level

of first gill openings plunlceti Waite 1910.

New Zealand, p. 89 ; also waitH
Thompson 1930, New Zealand, per-
haps a young stage of plunheti (see
discussion, p. 90-).

6. Head, to level of origin of pectoral fins, is only about 24-25 per cent
of total length ; caudal fin with distinct subterminal notch

*crepidater Bocage and Capello
1864, including *jonsonii Sae-
mundsson 1922. Eastern North
Atlantic, p. 91 ; also New Zea-
land, or represented there by a
close ally (see discussion, p. 93).
Head to level of origin of pectoral fins about 31 per cent of total

length, caudal fin without distinct subterminal notch

rossi Alcock 1898.
Indian Ocean, p. 94

Systematic position in the genus doubtful

fvscus Gilchrist and von Bonde
1924. Southern Africa, see discus-
sion, p. 95.

Genus SCYMNODON Boeage and Capello 1864

Seymnodon Bocage and Capello 1864, p. 263; 1866, p. 31, type species *S.

ringens Bocage and Capello 1864, p. 263; 1866, p. 32, PI. 1, fig. 1;

PI. 3, fig. 2A ; type locality off Portugal.
Generic Synonyms:
Ceniroplwrus in part, Giinther 1870, p. 423, for *Scymno(lnn ringens Bocage

and Capello 1864.
Zameus Jordan and Fowler 1903, type species Centrophorus squamulosu.t

Giinther 1877, Japan.
Not Ce7itrophorus Miiller and Henle 1837, type species *Sgualus granulosus

Bloch and Schneider 1801.
Generic characters. As in Centroscymnus (p. 85), except
with the upper teeth much longer midway along each side of
jaw than either toward the center of the mouth or toward its
outer corners (Fig. 13C). Lower teeth triangular, highest and
symmetrical along median sector of jaw, shorter but only weakly

BIGELOW AND SCHROEDER : SQUALOID SHARKS 97

oblique toward outer corners of mouth ; first and second dorsal
fin spines scarcely projecting beyond the skin ; dermal denticles
on sides of trunk scale-like, tridentate, the outer surface with
longitudinal ridges alone in some species, including the type but
also with cross ridges (Fig. 13, F, G) in one species {ohscurus
Vaillant 1888) that seems referable to Scyninodon by its denti-
tion.

Maximum recorded length 1100 mm. for ring ens (Bocage and
Capello 1866, p. 32).

Depth range. One of the four known representatives of this
genus has been reported from a depth as great as 1400-1435
meters (one specimen, Yaillant 1888, p. 68) ; a second, sqiiamu-
losus (Giinther 1887, p. 6), from 631 meters. A young specimen,
however (the only one known) of a third was found on the
beach at North Brighton, New Zealand (Archey 1921, p. 195).
No information in this respect is at hand for the type species
of the genus, so far known only from Portugal and from the
northern coast of Spain (Rey 1928, p. 457).

Remarks. Giinther (1870, p. 420) thought that "the passage
. . . from reclining to erect teeth in the lower ' ' jaw is too gradual
to justify the separation of Scyninodon from C entroscymnus ; or
the separation of either of these from CentropJiorus, for that
matter. But the difference in dentition between the species re-
ferred here to Scymnodo7i and those referred to C entroscymnus
(p. 88) is in fact so conspicuous that a glance at the mouth
is enough to place a given specimen in the one category or in
the other (provided it is properly referable to either), which
makes the retention of Scymnodon as a separate genus a matter
of convenience, even if nothing more.

The contrast is also interesting between the pointed or tri-
dentate denticles, with longitudinal ridges, on the sides of the
trunk on mature specimens of Scymnodo7i and the replacement
of denticles of this type by evenly rounded ones with concave
blades, in some of the species of Centroscymnus (p. 86). But
this difference can no longer be regarded as alternative between
the two genera, for the denticles are tridentate in the adults of
two of the species {*crepidater Bocage and Capello 1864, and
plunketi Waite 1909), the teeth of w^hich place them in Centro-
scymnus (p. 88). In any case choice of the denticles rather
than of the dentition as the primary distinction between Seym-

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BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

CO

w

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BIGELOW AND SCHROEDER : SQUALOID SHARKS 99

nodon and Ceniroscymnus, as by Garman (1913), bv Fowler
(1941), and by Bigelow and Schroeder (1948) would provide
no basis on which to refer a given specimen to the one genus
rather than to the other, unless it chanced to be full grown or
nearly so. AVhile the only illustration of one of the three known
species of Seym nodon {squamulosns Giinther 1877) show^s no
sign of fin spines, the artist seems to have overlooked them, for
Gunther (1887, p. 6, PI. 2, Fig. B) wrote of them as "very small,
scarcely projecting beyond the skin."

Species: Named species referable to Scymnodon as defined here
are : *S. ringens Bocage and Capello 1864, type species, offing of
Portugal; Centrophonis squamnlosus Giinther 1877, Japan; and
Ceniroscymnus ohscurus Vaillant 1888, off the coast of north-
western Africa,-^ in a haul from 1400-1435 meters. The New
Zealand shark, also, that was described by Archey (1921) as
Scymnodon sherwoodi has been retained in that genus by the
several authors (Whitley 1934, 1940; Fowler 1941; Big-elow,
Schroeder and Springer 1953 ; and Richardson and Garrick
1953), who have had occasion to refer to it. But we now learn
from Dr. J. A. F. Garrick" that the type (and only known)
specimen, which he has recently examined, shows no trace of
a spine in either of its dorsal fins, and that it is therefore one

Fig. 13. A, Scymnodon ringens, female, 893 mm., long, coast of Portugal,
ill British Museum. B, ventral view of anterior part of head of same.
C, upper and lower teeth of same, left hand side of mouth, x about 1,
midpoint of jaws marked bj' the dotted line. D, dermal denticles of same,
from side below first dorsal fin, x about 7. E, dermal denticles of same
from side of caudal peduncle x about 7. F, Scymnodon ohscuriis, dermal
denticle of type specimen, female, 590 mm. long, off northwest Africa
(p. 100), from side below first dorsal fin, x about 18, made available to us
through kindness of Dr. P. Budker. G, dermal denticle of same, after
Vaillant, 1888, PI. 2, Fig. 2. H, lower teeth of same, from drawing con-
tributed by Dr. P. Budker, x about 4. I, Scymnodon squamulosus, type
specimen, female, 658 mm. long, Japan, in British Museum, outline of
caudal fin from drawing contributed by Mr. Denys W. Tucker.

21 The locality as given by Vaillant (1888, p. 68), is "sur les c6tes du Soudan,"
a designation referring to the general region from southern Morocco to Senegal
(Folin, 1887. pp. 287-300).

22 Information contributed by Dr. Garrick in advance of publication.

100 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

of the Dalatiinae not of the Squalinae, as the sub-families are
defined here (p. 18).

We have pointed out elsewhere (Bigelow and Schroeder 1954,
p. 51) that the juvenile shark from the northwestern Atlantic,
on which Bigelow, Schroeder and Springer (1953, p. 233) based
their new species *Scymnodon melas, actually represented a
juvenile stage in the growth of *Centroscymnus coelolepis Bocage
and Capello 1864.

The dentition of ohscurus closely resembles that of *ringens,
as Dr. Paul Budker has kindly verified for us from examination
of the type specimen of ohscurus in the Paris Museum. Indeed,
Garman (1913, p. 208) has definitely rated ohscurus as a synonym
of *ringens. But a more abruptly truncate caudal fin, with
more definite subterminal notch and smaller pectorals in oh-
scurus (as pictured by Yaillant 1888, PI. 2, Fig. 2) than in
*ringens, and flank denticles with a double series of transverse
ridges between the more prominent longitudinal ridges (Fig. 13,
F, G; see also Vaillant 1888, PL 2, Fig. 2C) argue against this
union, hence ohscurus is retained here as a distinct species. It is
interesting, also, that in ohscurus some of the denticles on the
sides of the trunk of a given individual may be tridentate, while
others are not (Fig. 13, F, G).

The Japanese squamulosus (Giinther 1887, p. 5, PI. 2, Fig. B)
agrees with * ring ens in the shape of its denticles. But an exam-
ination of the type specimen in the British Museum by Mr.
Denys W. Tucker (to whom we are greatly indebted), shows
its gill openings as only about 50 per cent as long, relatively, as
those of *ringens; its snout as longer relatively and more
pointed (length from tip of snout to mouth about as great as
distance between inner ends of preoral clefts) ; and both its sec-
ond dorsal, its pectoral and ventral fins, also its caudal (Fig.
13, I), as differing so conspicuously from those of *ringens, that
the two species are clearly distinct. In all these respects, in-
deed, squamulosus so nearly resembles ohscurus (as pictured by
Vaillant 1888, PI. 2, Fig. 2) that these two species might easily
be confused. But the denticles on the sides of the trunk of
squamulosus (as pictured by Giinther 1887, PI. 2, fig. B) show
no trace of the transverse ridges that mark the denticles of
ohscurus (Fig. 13, F, G).

BIQELOW AND SCHROEDER : SQUALOID SHARKS 101

Key to Species of Scymnodon

1. Lower outline of caudal fin nearly continuously rounded, -with sub-
terminal notch only faintly marked, and without definite lower-anterior
lobe (Fig. 13 A) ; pectoral fin, when laid back, reaching very nearly
to a perpendicular at base of first dorsal fin spine

*ringens Bocage and Capello 1864,
eastern North Atlantic, p. 99
Lower outline of caudal fin with well-marked subterminal notch and
prominent lower anterior lobe (Fig. 13, I) ; pectoral fin, when laid back,
falls short of a perpendicular at base of first dorsal fin spine by a
distance at least 50 per cent as long as snout anterior to eyes 2

2. Outer surface of dermal denticles on sides of trunk with weak trans-
verse ridges in addition to the three longitudinal ridges (Fig. 13, F,
G) ohscurus Vaillant 1888, off coast of

Northwestern Africa, p. 100
Outer surface of dermal denticles on sides of trunk without trans-
verse ridges squamulosus Giinther 1877, Japan, p. 100

Genus DeaNIA Jordan and Snyder 1902

Deania Jordan and Snyder 1902, p. 80, type species, *D. eglantina Jordan

and Snyder, Japan.
Generic Synonyms:

Acanthidium in part, Lowe, 1839, p. 92, for *A. calceum Lowe, Madeira.
Centrophorus in part, Lowe, 1843, p. 93, and subsequent authors, for C.

calceus Lowe 1843, equals * Acanthidium calceum Lowe 1839.
Centrophorus in part, Bocage and Capello 1864, 1866, for C. crepidalius

Bocage and Capello, evidently equals *Acanthidiuvi calceum Lowe 1839.
Scymnodon in part, Goode and Bean 1895, by error, their PI. 4, Fig. 12,

labelled ' ' S. ringens Bocage and Capello ' ' in the explanation of

plates being a copy of the illustration by Bocage and Capello (1866,

PI. 2, Fig. 1) of Centrophorus crepidalhus Bocage and Capello 1864,

equals Acanthidium calceum Lowe 1839.
Nasisqualus Smith and Eadcliffe 1912, p. 681, type species *N . profundorum

Smith and Eadcliffe, Philippines.
Acnnthidium Garman 1913, p. 215; type designated as *A. calceum Lowe

1839; but incorrectly so, because *A. pusillum Lowe 1839 (which is a

species of Etmopterus Eafinesque 1810, see below) had previously been

designated as the type of Acanthidium by Jordan and Evermann

(1896, p. 55).
Daeniops Whitley 1932, p. 36, type species Acanthidium, quadrispinosum

McCulloch 1915, Australia.

102 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Not Centrophonis Miiller and Henle 1837, type species *Sqiuilus granulosus

Bloeh and Schneider 1801.
Not Acanthidium Lowe 1839, type species *A. pu^illum Lowe as designated

by Jordan and Everraann 1896, p. 55, and by Jordan 1919, p. 195,

making Acanthidium a synonym of Etmopterus Eafinesque 1810 (sec

above).
Not Scymnodon Bocage and Capello 1864, 1866, type species *S. ringenn

Bocage and Capello 1864.

Generic characters. Inner corner of pectoral fins rounded in
most species, perhaps rectangular in some, but not extended
(p. 103) ; snout in front of mouth longer than from mouth to
level of origin of pectoral fins; preoral clefts only very slightly
expanded inwardly ; dermal denticles on sides of trunk high,
pitchfork shaped ; teeth smooth-edged in all known species ; other
characters as in CentropJiorus (p. 64).

Maximum recorded lengths, 1138 mm. {qnadrispinosa Mc-
(.^ulloch 1915, p. 100) ; 1070 mm. {kaikourae Thompson 1930, as
"calceus") ; and 1060 mm. {calcea, Vaillant 1888, p. 71).

Depth range. The depths of capture so far recorded for
Deaiiia (mostly, at least, in the trawl, hence subject to the un-
certainty emphasized on p. 5) are: 605-1431 meters for calcea
(Roule 1919, p. 119) ; 715-1785 meters for *profundorum (Smith
and Radcliffe 1912, p. 683) ; 190-366 meters for *eglantina (Smith
1949, p. 58) ; and 238-823 meters for qnadrispinosa (McCulloeh
1915, p. 100; Whitley 1940, p. 147). While definite information
as to depth is wanting for kaikourae, it was reported from "deep
water" (Thompson 1930, p. 276, as "calceus"). And the Jap-
anese records for *acicidata (Garman) 1906, *rostrata Garman
1906, and hystricosa (Garman) 1906 which we here refer to
*eglantina (p. 104) were from the deep line-fishery in Sagami
Ba3^ Evidently Deania can fairly be termed a deep-water genus.

Remarks. The chief respects in which the various species that
have been grouped together as Deania (or as one or other of its
synonyms) difiler from Centrophonis are: a much longer snout,
and pitchfork-shaped denticles on the sides of the body. Most
of the species of Deania are set further apart from Centrophonis
by the rounded shape of the inner corner of their pectoral fins.
But the two genera intergrade in this respect, the inner corner
of the pectorals being quadrate, both in one member of Centro-
phonis ifoliaceus Giinther 1877), as noted above (p. 66), and

BIGELOW AND SCHROEDER : SQUALOID SHARKS 103

also in one member of Deania, if Gilchrist's (1922, PI. 7, Fig. 2)
illustration of his D. natalense represents the latter correctly.^^

Deania differs from Centroscymnns and from Scymnodon in a
much longer snout; in a longer exposed portion of the second
dorsal fin spine ; and in broader upper teeth. It differs further
from Scymnodon in that the teeth in the central part of its upper
jaw are nearly or quite as long as those midway out along either
side. It had appeared also until recently that its high, pitch-
fork-shaped denticles were equally diagnostic of Deania as con-
trasted with Centrophorus and with the C entroscymnus-Scymno-
don group. But Garriek (1955, p. 234, Fig. 2) has found that
the denticles on the sides of the trunk of young specimens of
Centroscymnus ivaitei (Thompson) 1930 of New Zealand (p. 90)
approach the pitchfork-like shape that was formerly thought to
be diagnostic of the denticles of Deania.

The cusps of the upper teeth of Deania are sharp-tipped,
rather narrowly triangular, with concave margins, on broad
bases ;'■* erect along the central part of the jaw, but increasingly
oblique toward the outer corners of the mouth.

The lower teeth are very strongly oblique all along either side,
and the median tooth is not symmetrical in the female of any
species for which the condition in this respect is known. In
*calcea (Lowe) 1839 (the oldest species), the lowers are of the
same shape in the males as in the females, as illustrated by a
male of 848 mm. and a female of 1037 mm. that we have seen
from southwest of the Faroes.'^ Similarly, the lowers are de-
scribed and pictured as stronglj^ oblique in the male in *pro-
fundorum (Smith and Radcliffe 1912, p. 681, PI. 53), which
we can verify from our examination of the type specimen. But
the lowers are described by McCulloch (1915, p. 101) as much
less strongly oblique in the males of quadrispinosa, from
Australia, than in the females, which is also the case in the
Japanese species-complex that we unite here (p. 104) under
the name *eglantina Jordan and Snyder 1902. Thus the lowers
(including the median tooth) of the female pictured by Garman

23 The pectoral fins are not mentioned in Gilchrist's (1922, p. 49) description
of his natalense.

2* Our re-examinatlon of the type specimen of *Deania eglantina (type of the
genus) has shown that Jordan and Snyder's (1902, p. 81) description of the
teeth as with "small basal cusp" is not correct.

25 We are indebted to Dr. J. R. Pfaff, of the University Zoological Museum,
Copenhagen, for the opportunity of seeing these specimens.

104 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

(1913, PI. 11, Fig. 7) as Acanthidmm hystricosum^^ are shown
as strongly oblique, whereas those of the adult male figured
by him (Garman 1913, PI. 12, Fig. 2) as *Acanthidium acicula-
tum (which we have examined) are erect in the front of the
mouth, with the median tooth symmetrical. And the reserve rows
of lower teeth of an immature male, 682 mm. long, in the
Museum of Comparative Zoology, are similarly nearly erect along
the center of the jaws (the functional row is badly worn). The
situation in this respect is not known for natalense or for
kaikourae.

It is a matter of common knowledge that sexual dimorphism
of this sort is common among batoid elasmobranchs, but it seems
not to have been reported previously among sharks.

Species: Named species referable to Deania: are * Acantkidium
calceum Lowe, 1839, eastern Atlantic; * Deania eglantina Jordan
and Snyder 1902, type species, Japan ; * Acanthidium, acicMlatum
Garman 1906, A. hystricosum Garman 1906, and *A. rostratum
Garman 1906, all from Japan ; *Nasisqualus profimdorum Smith
and Radcliffe 1912, Philippines; Acanthidium quadrispinosum
McCulloch 1915, Australia; Acanthidium natalense Gilchrist
1922, Natal, southeastern Africa; and Centrophorus kaikourae
Whitley 1934, New Zealand, which was originally reported and
pictured by Thompson (1930, p. 275, PI. 42 as Centrophorus
calceus Lowe).

Our re-examination of the type specimens of *eglantina (a
female of 300 mm.), *aciculata (a male of 890 mm.) and *ro-
strata (a female of 873 mm.) has revealed no differences between
them in proportional dimensions (see below) greater than may
reasonably be credited to the difference between them in stage of
growth, or to the variability that has already been reported by
Regan (1908, p. 52) for the allied *calcea of the Atlantic.
Cases in point are the somewhat longer head and caudal fin, but
shorter interdorsal space of the * eglantina. And while the type
specimen of the fourth member of this group {hystricosa) is no
longer to be found, Carman's (1913, PI. 11, Figs. 5-8) beautiful
illustrations of it, by the well-known zoological artist E. N.
Fischer, do not suggest anything to set it apart either. Hence,
all deanias yet reported from Japan are referred here to the
one species * eglantina Jordan and Snyder 1902.

26 This specimen is no longer to be found.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 105

Proportional dimensions in per cent of total length. A. *Deania
eglantina, female, 300 mm., type, U.S.N.M. No. 49524; B.
*D. rostrata, female, 873 mm., type, M.C.Z. No. 1047; C. *D.
aciculata, male 890 mm., type, M.C.Z. No. 1128; all from Japan.

A

B

C

Trunk at origin of pectoral: breadth

9.3

7.5

6.7

height

7.7

8.0

6.7

Snout in front of: outer nostrils

5.3

5.3

4.0

mouth opening

15.3

13.7

12.1

Eye: horizontal diameter

5.3

3.6

4.7

Mouth: breadth

7.7

5.6

6.4

Nostrils: distance between inner ends

4.5

3.4

3.4

Preoral clefts: distance between inner ends

4.8

3.4

3.7

Gill opening lengths: first

2.0

—

1.7

third

2.0

2.5

1.7

fifth

2.0

2.1

1.8

First dorsal fin: vertical height

3.8

3.5

2.9

length of l)ase from spine origin

9.5

9.7

11.8

Second dorsal fin: vertical height

4.7

5.5

4.8

length of base from spine origin

9.2

10.0

10.0

Caudal fin: upper margin

20.0

16.0

16.8

lower anterior margin

11.6

10.8

9.8

Pectoral fin : width

6.2

6.1

6.1

length of anterior margin

9.3

9.0

8.9

Distance from snout to : 1st dorsal spine

42.3

39.9

40.2

2nd dorsal spine

68.2

70.5

70.5

upper caudal

80.0

84.0

83.2

pectoral

28.7

24.0

23.2

pelvics

61.6

64.7

62.6

Interspace hetiveen: origins of first and

second dorsal spines, at base

24.6

30.5

30.7

second dorsal and caudal

3.8

3.2

3.4

pelvics and caudal

8.9

10.3

10.1

Distance from origin to origin of :

pectorals and pelvics

33.3

40.8

39.3

pelvics and caudal

13.3

13.4

14.4

Teeth, ■number of series: uppers

28

34

29

lowers

13-1-14

?

29

The Japanese *eglantina (inclnding Garman's supposedly dis-
tinct species, as just noted), is so closely allied to *Deania calcea
of the eastern Atlantic that Regan (1908, p. 51) has already
relegated it unequivocally to the synonymy of * calcea, and

106 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

*aciculata doubtfully so. But it seems to us wiser to retain
*eglantina as a separate species, at least until a large number of
specimens of the Japanese and North Atlantic populations have
been studied, for while the Japanese specimens listed above agree
very closel}' indeed Avith the two *calcea we have seen, in fin
characters and in bodily proportions in general, the lower teeth
are as oblique in the males of *calcea as in the females, but are
more erect in mature males than in females in *cglantina, as
noted above (p. 103). Though this last difference would not be
an aid to identification unless the species in hand chanced to be
well-grown males, it deserves taxonomic recognition of some
sort. And the ([uestion as to whether the North Atlantic and
North Pacific populations of this general type have diverged
significantly since they became isolated geographically, one from
the other, is one too interesting to be answered offhand.

The * calcca-* eglantina group of the Northern Hemisphere
has close counterparts, in the Southern Hemisphere, in quadri-
spinosa of Australia ; also in a shark reported and pictured from
southern African waters by Smith (1949, p. 58, Fig. 49) as
'*eglantina, which certainly resembles the type specimen of *eg-
lantina very closely. But the eventual decision as to the identity
of either or both of these with ^eglaniina of Japan must await
the comparison of specimens from the different ocean areas.

In the * calcea-* eglantina-quadrispinosa group (however many
species this may eventually prove to represent), the pectoral
fins, when laid back, fall considerably short of a vertical at the
point of emergence from the skin of the first dorsal fin spine.
But the pectorals of *profundorum from the Philippines are
pictured as reaching rearward to abreast of the first dorsal spine
(Smith and Radclift'e, 1912, PI. 53), and our own examination
of the type specimen has shown this to be correct. As this is a
rather conspicuous difference, * prof undornm may be accepted
as a good species.

The original illustration of natale7ise (Gilchrist 1922, PI. 7,
Pig. 2) of southern African waters, shows the pectorals not
only as reaching rearward beyond the first dorsal spine, but also
with the inner corner quadrate, instead of rounded as they are
in other known representatives of the genus Deania. But Smith's
(1949, p. 58) allocation of it (with quadrispinosa) to the syn-
onymy of *eglanihia — perhaps from examination of the type

BIQELOW AND SCHROEDER : SQUALOID SHARKS 107

which Barnard (1925, p. 51) has reported as being in the collec-
tion of the Government Marine Surve}- of South Africa —
suggests that the artist's representation of natalensc may not
have been correct. Hence its status remains in doubt, pending
further information.

Finally, kaikourae (falling with the * calcea-* eglantina group
in length of pectoral fin) differs from all other known members
of Deania in a first dorsal fin base about 1.6 times as long as the
second (each measured from the point of emergence of the
respective fin spine from the skin), contrasted with 0.9-1.2
times for the *calcea-*eglantina-quadrispinosa group, for *pro-
fundorum, and for natalcnse. Thompson (1930, p. 276) indeed,
has already called attention to this feature in his original ac-
count of kaikourae (as "CentropJiorus calceus").

To sum up, the nine representatives of the genus Deania that
have been named from various seas may represent as few as four
species, *calcea in the North Atlantic, '*e.glantiiia in the North
and South Pacific and southern Indian Ocean, kaikourae from
New Zealand, and *profundorum in Philippine waters. Or they
may represent as many as six, if quadrispinosa from Australia,
and the South African form reported by Gilchrist 1922 as 7iai-
alense, but by Smith (1949) as '*eglantina, should finally prove
to be separable, not only from *cglantina of the North Pacific,
but one from the other as well.

Key to Species of Deania

1. Pectorals, when laid back, falling considerably short of a perpendicu

lar at point of emergence of first dorsal fiu spine from the skin 2

Pectorals, when laid back, reach at least as far as a perpendicular at
point of emergence of first dorsal fin spine 4

2. First dorsal fin, from point of emergence of spine to rear end of base
is 1.5-1.6 times as long as second dorsal, similarly measured

Icaikourae Whitley 1934, Australia, p. 107
First dorsal fin, from point of emergence of fin spine to rear end of
base of fin, is only 0.9-1.2 times as long as second dorsal fin, similarly
measured 3

3. Lower teeth as strongly oblique in males .is in females

*calcea Lowe 1939.

Northeastern Atlantic, including Mediterranean, p. 105

Lower teeth more erect in males than in females *eglantina

108

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Figure 14

BIGELOW AND SCHROEDER : SQUALOID SHARKS 109

Jordan and Snyder 1902 (incl. *aci-
culata Garman 1906, *rostrata Gar-
man 1906, and hystricosa Garman
1906), Japan, and reported from
southern Africa (p. 106) ; also
quadrispinosa Gilchrist 1922, Au-
stralia, p. 106
4. Inner corner of pectoral fins rounded; first dorsal fin only about

70 per cent as high vertically as second dorsal fin

*profundorwm Smith and

Eadcliffe 1912, Philippines, p. 106

Inner corner of pectoral fins quadrate; first dorsal fin about as high

vertically as second dorsal fin natalense Gilchrist 1922.

Off Natal coast, southern Africa
(but see discussion), p. 107

Subfamily DALATIINAE

Subfamily characters. Squalidae lacking a second dorsal fin
spine (lacking a first spine also in most cases) ; preoral clefts
lacking in most but present in one of the known genera (Scym-
iwdalatias, p. 124) ; dermal denticles either truncate with hollow
crown, or with a single cusp or spine ; teeth with only one cusp,
the uppers narrow, awl shaped or lancet shaped ; the lowers much
broader, with quadrate base, each overlapping the next outward ;
caudal axis raised in some but not in others ; the caudal fin
varying widely in shape accordingly (Fig. 14), its tip more or
less evidently truncate ; sides of body, anterior to cloaca, with-
out longitudinal ridges.

Fig. 14. Outlines of caudal fins of representative genera, adjusted to
equal lengths along upper margin, to show relative breadth of fin above
axis and below ; also degree to which caudal axis is raised. A, Dalatias
UcJm, male, 1114 mm. long, Japan, Mus. Comp. Zool. No. 1116. B, Somnio-
diis microcephalus, male, 1334 mm. long, Massachusetts Bay, Mus. Comp.
Zool. No. 39609. C, Isistius brasilietisis, male, 383 mm. long, western North
Atlantic north of the Bahamas, Mus. Comp. Zool. No. 36039. D, Squaliolus
laticaudus, same specimen as in Fig. 16A. E, Echinorhinus bruous, male,
1650 mm. long, Mauritanian coast, northwest Africa, Mus. Comp. Zool. No.
39633. F, Lanina ditropis, type specimen, male, 2085 mm. long, off La Jolla,
California, Mus. Comp. Zool. No. 36471, the vertebral axis indicated by the
broken line.

110

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Figure 15

BTGELOW AND SCHROEDER : SQUALOID SHARKS 111

Genera. The shark species referable to this admittedly "arti-
ficial" group have been distributed among the following named
genera: (1) DaJatias Rafinesque 1810, type species D. sparo-
phagus Rafinesque, equals *Squalus licha Bonnaterre 1788; (2)
Somniosus Lesueur 1818, type species iS^. hrevipmna Lesueur,
equals *Squalus microcephalus Bloch and Schneider 1801; (3)
Euprotomicrus Gill 1864, type species Scymnus {Laemargiis)
labordii Miiller and Henle 1841, equals *Scymnus hispinatus
Quoy and Gaimard 1824; (4) Isis-tius Gill 1864, type species
*Scymnus brasiliejisis Quoy and Gaimard 1824; (5) Heteroscyni-
nus Tanaka 1912, type species *n. longus Tanaka; (6) Squalio-
lus Smith and Radcliffe 1912, type species *S. longicaudus Smith
and Radcliffe; (7) Heteroscymnoides Fowler 1934, type species
*H. marleyi Fowler; (8) Pseudoscymmis Herre 1935, type
species P. hoshuensis Herre; and (9) Scymnodalafias Garrick
1956, type species Scymnodon sherwoodi Archey 1921. There
seems nothing, however, to set Pseudoscymnus apart from Dal-
atias (Bigelow and Schroeder 1948, pp. 501, 502). And Hubbs
and McHugh's (1951, p. 164, Footnote) suggestion that Hetero-
scymnus should be reduced "perhaps to the synonymy of
Somniosus," is supported by our examination of the teeth (PL
3) and denticles (Fig. 15B) of the one known species, *//.
longus Tanaka 1912, from Japan. For further discussion of it
see page 121.

Fig. l.j. A, Somniosus longus, type specimen, female, 13G0 luni. luny,
.lapan, al'ter Tanaka, the .shapes of the fins somewhat emended to accord
with a Japanese male, 1010 mm. long, Miis. Comp. Zool. No. 396r)0, kindly
(ontiibuted by Dr. Tokiharu Abe. B, dermal denticles of latter, from side
below first dorsal fin, x about 25. C, side view of same. D, Somniosus
rostratus, lower tooth of Mediten-anean female, 820 mm. long, after
Canestrini, ISfi-l, PI. 2, Fig. 4. E, lower teeth near center of jaw, of
Portuguese shark described by Capello 1870 under the name " rostratus'',
traced from a photograph contributed by Dr. A. M. Eamalho. F, Hetero-
scymnoides marleyi, type specimen (see Fig. 16E), upper and lower teeth
at center of mouth, x about 12. G, dermal denticles of same, from side
bv-low first dorsal fin, x 36. H, Squaliolus laticaudus, same specimen as in
Figure IGA, denticles on side below first dorsal fin, x about 45. I, Dalatias
licha, same specimen as in Figure 14 A, a sector of the exposed part of
upper and lower lips, x about 4.

112 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

In short, the dozen or so known species of Dalatiinae that
appear to deserve separate recognition seem referable to 6 or 7
genera as defined by the positions and relative sizes of the first
and second dorsal fins, presence or absence of a spine on the first
dorsal fin, nature of the teeth, shape of the dermal denticles on
the sides of the body, and degree to which the portion of the
caudal fin above the axis is developed relative to the portion
below the axis. This last matter is one of especial interest, as
already stressed by Hubbs and McHugh (1951, p. 164) in their
key to genera, for in the genera in which the fin is the most
widely expanded above the axis (Fig. 14), the latter is raised
only very slightly above the main axis of the shark's body (if
at all). In other words, one of the evolutionary trends of the
group has been a tendency for the tail to lose the heterocercal
nature so characteristic for it among sharks in general. Cau-
dal fins even more lunate in form (Fig. 14F) are, it is true,
well known characteristics of the isurid, basking, and whale
sharks. But in all of these the caudal axis is steeply raised ; they
are, in a word, as truly heterocercal as any other shark, more
noticeably so, indeed, than many, such as the erectolobids,
scyliorhinids and some of the triakids.

Key to Genera of Dalatiinae

1. Lips with a complex series of fringed cross folds (Fig. 15, I) ; margins

of lower teeth regularly serrate Dalaiias Eafinesque 1810. p. 113

Lips smooth, or nearly so; margins of lower teeth smooth, or only
faintly and partially serrate at most 2

2. First dorsal fin with a spine, the tip either exposed or buried in the

skin Squaliolus Smith and Radcliffe 1912. p. 128

First dorsal fin with no trace of a spine 3

3. Preoral clefts present ; upper margin of caudal fin about twice as long

as lower anterior margin Scymnodalatias Garrick 1956. p. 124

No preoral clefts ; upper margin of caudal fin not more than 1.5-1.7
times as long as lower anterior margin, and relatively shorter than
this in most 4

4. Eear end of base of first dorsal fin is over or posterior to origin of
pelvic fins ; interspace between first and second dorsal fins is shorter
than between second dorsal fin and origin of upper margin of caudal ;
lower teeth erect, symmetrical Isistiiis Gill 1864. p. 123

BIGELOW AND SCHROEDER : SQUALOID SHARKS 113

Rear end of base of first dorsal fin is clearly anterior to origin of
pelvics; interspace between first and second dorsal fins is at least as
long as between second dorsal fin and origin of upper side of caudal ;
lower teeth asymmetrical, the cusp more or less strongly oblique,
outward 5

5. Origin of first dorsal fin is about over axil of pectorals

Hetcroseymnoidcs Fowler 1934. p. 182
Origin of first dorsal fin is posterior to tips of pectorals when these are
laid back 6

6. Base of first dorsal fin about as long as base of second dorsal ; eye
only about 20-25 per cent as long as snout in front of mouth

Somniosus Lesueur 1818. p. 115
Base of first dorsal fin only about 25-33 per cent as long as base of
second dorsal ; eye more than nO per cent as long as snout in front of
mouth Euprotomicrvs Gill 1864. p. 126

Genus DaLATIAS Rafinesque 1810

Dalatias Rafinesque 1810,^ p. 10; type species D. sparophagus Rafinesque,
Mediterranean, equals *Squalus licha Bonnaterre 1788, designated by
Jordan, Tanaka and Snyder 1913. For generic synonyms and refer-
ences, see Bigelow and Schroedcr 1948, p. 501.
Geyieric characters. Dalatiinae with blade-like lower teeth, the
cusp erect, triangular, with serrate edges, the base quadrate
with a conspicuous notch on the outer edge marking the transi-
tion from cusp to base. Snout in front of mouth much shorter
than from mouth to level of first gill openings ; longest gill open-
ings about 8-11 per cent as long as head to origin of pectorals
(specimens measured) ; lips only moderately expanded at corners
of mouth, and without special cartilages, but conspicuously thick
and fleshy inward along each jaw and close-set with a complex
series of cross folds with finely fringed edges (Fig. 15, I) ; fur-
row from corner of mouth reaching rearward about 30 per
cent of distance towards first gill openings ; no preoral clefts, but
with voluminous subdermal pouches extending forward from
the oral pockets (Fig. IE) ; caudal fin of the shape shown in
Fig. 14A, its axis rather steeply raised ; origin of first dorsal fin
much nearer to rear edge of pectorals when these are laid back
than to origin of pelvics ; second dorsal fin about 1.3 times as
long as first dorsal at base ; no spine in either of the dorsal
fins. Denticles on sides of body thick, quadrate, outer surface
with three weak ridges united rearward in a point that varies

114 BULLETIN : MUSEUM OP COMPAEATIVE ZOOLOGY

in length and in aeuteness from denticle to denticle; denticles
on lower side of snout overlapping, ovate, without marginal
teeth (Bigelow and Sehroeder 1948, Fig. 96, D, E).

Greatest recorded length, 1820 mm. (Dumeril 1865, p. 452).

Depth range. Definite depths of capture that we have found
recorded for Dalatias in one part of the oceans or another have
ranged from about 90 meters down to 600 meters. Probably it
is more numerous in the sboaler part of its vertical range, else
fishermen would hardly be as familiar with it as they have long
been in the eastern side of the North Atlantic, from northwestern
Africa and the Canaries to the Irish slope ; also in the Mediter-
ranean.

Remarks. The serrate lower teeth of Dalatias, and the shape
of its caudal fin, with its conspicuously fleshy and complexly
sculptured lips, set it so sharply apart that there is no danger of
confusing it with any other genus of Dalatiinae.

Species. It is probable that all the reports for Dalatias that
have appeared in scientific literature during the past century
and a half have been based on the single species *D. licha
(Bonnaterre) 1788, type of the genus, which was originally de-
scribed from "Cap Breton" on the coast of Brittany, by Brous-
sonet (1780, p. 677), as "La Liche." In the eastern side of
the Atlantic, Hicha is now known to be generally distributed
from tropical West Africa (Rio de Oro), Morocco, the western
Mediterranean, the Canaries, and Madeira to the Irish Atlantic
slope and to the fishing grounds west of Ireland. Contrasting
with this widespread occurrence, and with its local abundance
in the Mediterranean and near Ireland, there have been onlv
two pul_)lished records of its capture in the western Atlantic,
the one for a female of about 5 feet (about 1550 mm.), taken
on Georges Bank, August 19, 1937 (Nichols and Firth 1939, p.
85; Bigelow and Sehroeder 1948, p. 502) ; the other for an 845
mm. female from the northern part of the Giilf of Mexico (Big-
elow, Sehroeder and Springer 1955, p. 10). But the recent cap-
ture by "Oregon" of a female only about 380 mm. long (U. S.
Nat. Mus. No. 157844) in the northern side of the Gulf, sug-
gests that the latter may harbor a resident population.

*D. licha has also been reported under its o-\vn name by Smith
(1949, p. 56) from southern Africa, whence it had previoush
been recorded as a separate species [D. hrevipinnis Smitli

BIGELOW AND SCHROEDER : SQUALOID SHARKS 115

1936, p. 1); from New Zealand; from Australia; also from
Japan both as JicJia and as americanus Gmelin 1789 (an obvious
synonym for h'cha). Identification as licha in the case of the
Japanese form is corroborated by our own comparison of a
Japanese specimen with the specimens we have seen from Georges
Bank and from the Gulf of Mexico. (For the list of New Zea-
land, Australian and Japanese references for *licha, see Bigelow
and Schroeder 1948, p. 508.) Nor do we find anything in
Herre's (1935, p. 124) account of the Japanese shark for which
he proposed the new generic and specific names Pseud oscymnus
hohuensis to suggest that it differs in any way from *Z). licha.
And this applies equally to McCulloch's (1914, p. 81, PI. 14)
excellent account and illustration of the Australian shark that
he described as "licha" but which Whitley (1931, p. 310; 1940,
p. 151) rechristened phillippsi, although without including any
supporting evidence to justify the new specific name.

AVith the known range of *Z>. licha so extensive, it seems
astonishing that the genus has not been reported as yet from
the eastern side of the Pacific, south or north, or from the
southern Atlantic, though it is known from Algoa Bay, south-
eastern Africa.

Genus SOMNIOSUS Lesueur 1818

Somniosiis Lesueur 1818, p. 222; type species S. brevipinna Lesueur, Massa-
rhusetts, equals *Squalus TuicrocepJialus Bloch and Schneider 1801.

For generic synonyms, see the list given by Bigelow and Schroeder 1948,
p. 514, with the addition of Iletcroscymnus Tanaka 1912, type species 77.
longus Tanaka, Japan (see p. 121).

Generic characters. Dalatiinae, without fin spines ; teeth smooth
edged, the lowers with the cusp directed so strongly outward in
some species that the inner edges function as a nearly unbroken
cutting edge parallel with the jaw, but more nearly erect in
others (PI. 3) ; snout in front of mouth at least not longer than
from mouth to level of first gill openings; longest gill openings
about 9-11 per cent as long as head to origin of pectorals on
specimens measured ; lips not conspicuously thick, their surface
smooth ; furrows from corners of mouth reaching rearward only
about 35-40 per cent of the distance toward first gill openings;
no preoral clefts but with voluminous preoral pouches extend-

116 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

ing forward from oral pockets; caudal fin with upper division
widely expanded, as well as the lower (Fig. 14B) ; upper margin
about 1.5 times as long as lower anterior margin. Caudal axis
moderately raised. Origin of first dorsal fin nearer to rear edge
of pectorals (when these are laid back) than to origin of pelvics;
second dorsal fin not larger than the first dorsal ; interspace
between first and second dorsal fins 1.7-3 times as long as be-
tween second dorsal and caudal ; denticles on sides of body coni-
cal to thorn-like, curved rearward, differing widely in size from
species to species, and in degree of elevation from the skin.

Size. The different species of Somniosus differ widely in
size. Thus rostratus is not known to grow longer than about one
meter. But microcephalus averages 8-14 feet (2.4-4.3 meters) at
maturity, with specimens of 16-18 feet (4.9-5.5 meters) taken
occasionally, and one of 21 feet (6.4 meters) definitely recorded
(see Bigelow and Schroeder 1948, p. 519 for further informa-
tion).

Depth range. The best known member of the genus {*micro-
cephalus, p. 117) has long been known to occupy a wide depth
range, coming right up to the ice in the northern part of its
range (Greenland, Labrador) in winter, but in summer most
often caught at 180-550 meters, with one recorded from 1207
meters. (For further details see Bigelow and Schroeder 1948,
p. 521.) And the depth range is, presumably, about the same
for its representative in the North Pacific {pacificus, p. 121),
though precise information is lacking. The only definite depth
record we find for S. rostratus of the eastern Atlantic, said by
Tortonese (1937-1938, p. 75) to occur off Portugal in shoal
water as well as deep, is about 1000 meters, near Madeira, by
line fishermen's report (Maul, 1955, p. 7). And the only
information at hand in this respect for *S. longus of Japan
(p. 121) is that the s*pecimens so far seen Avere the product of
the deep hook and line fishery in Sagami Bay, Tanaka 1912a,
p. 104).

Remarks. The combination of a very short snout with a caudal
of the shape shown in Figure 14B, a first dorsal fin originating
near the mid-length of the trunk, a second dorsal not larger
than the first, and thorn-like denticles, with lower teeth more or
less strongly oblique, and razor sharp, makes the recognition of

BIGELOW AND SCHROEDER : SQUALOID SHARKS 117

a Somniosus easy, even as one is dumped on deck from the
trawl.

The caudal fin in Somniosus has a distinct subterminal notch
in the specimens we have seen from the Gulf of Maine, from
Iceland, and from Japan ; indeed it has been so pictured for
rostratus by Burkhardt (1900, p. 564, Fig. 3) and by Helbing
(1904, PI. 9, fig. 1) ; for *microcephalus by Helbing (1904, PI.
9, fig. 2, as "horealis") ; by Garman (1913, PI. 15, fig. 4), and
by Bigelow and Schroeder (1948, Fig. 100) ; by Tanaka (1912a,
PI. 26) for Hongiis. Since it is similarly notched on the type
specimen of *>S^. pacificus Bigelow and Schroeder 1944, from Japan
(pictured by Garman 1913, PI. 15, fig. 1, under the name
"hrevipinna"), we suspect that Tanaka 's (1911, PI. 11, fig. 32)
earlier illustration of it for that species (under the name
microcephalus Bloch and Schneider 1801) as without a notch,
represents the result either of wear, or of mutilation.

More significant from the taxonomic standpoint is the wide
expansion of the fin above the caudal axis, and its sublunate
shape (Fig. 14B).

One species {rostratus of the Mediterranean and adjacent
Atlantic) is brilliantly luminescent. For a first hand account of
its luminescence, and of its light-producing organs, see Burk-
hardt 1900, p. 560.

Species. The following named species (listed here in chrono-
logical order) fall in Somniosus :

*Squali(s microcephalus Bloch and Schneider 1801, boreal-
subarctic latitudes in the eastern Atlantic ; *Somniosus brevi-
pinna Lesueur 1818, corresponding zone in the western North
Atlantic; Scymnus rostratus Risso 1826, Mediterranean and
Madeira; *Hetero scymnus longus Tanaka 1912, Japan; Somnio-
sus antarcticus Whitley 1939, Macquarie Island, Subantarctica,
south of Australia; * Somniosus pacijlcus Bigelow and Schroe-
der 1944, Japan, probably also northeastern Asiatic coast, and
American coast from Bering Strait southward to Puget Sound,
occasionally to southern California.

Our own comparison of a 1334 mm. male, taken off the coast
of Massachusetts, with a 1564 mm. male from Iceland corrobo-
rates the view, now generally held, that the Greenland sharks of
the subarctic on the two sides of the Atlantic (including the
White Sea) and of the neighboring parts of the Arctic Ocean

118

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

belong to a single species. How closely the population of *S.
microcephalus in the two sides of the North Atlantic agree in
proportional dimensions appears from the following tabulation :
A, for a male, 1334 mm. long taken near Boston Lightship in 32
fathoms (Mus. Comp. Zool. No. 39609) and B, for a second
male, 1564 mm. long, taken northwest of Iceland, Lat. 66°48'N.,
Long. 25°10'W., in 180 fathoms, October 1952 (Mus. Comp. Zool.
No. 37826).

Proportional dimensions in per cent of total length

Snout length in front of : outer nostrils

eye
mouth
Eye: horizontal diameter
Mouth: breadth

Nostrii.s: di.stance between inner ends
cm opening lengths: first

second
third
fourth
fifth
First dorsal fin: vertical height
length of base
Second dorsal fin : vertical height

length of base
Caudal fin: upper margin

lower anterior margin
Pectoral fin: outer margin
inner margin
greatest width
Distance from snout to : first dorsal

second dorsal
upper caudal
pectorals
pelvics
interspace between: first and second dorsals
second dorsal and upper caudal
pelvics and lower caudal
Distance between origins of: pectorals and pelvics
pelvics and subcaudal

Teeth :

A

3.1

B

2.4

7.3

6.4

10.3

9.3

2.2

2.4

6.7

7.2

3.9

3.4

1.8

2.0

2.0

2.2

2.0

2.2

2.0

2.2

1.8

2.0

3.4

3.5

7.6

8.4

2.9

2.6

6.6

5.8

20.5

20.6

14.9

15.0

12.1

13.2

6.0

5.6

6.5

6.2

41.2

38.0

65.3

64.9

79.5

79.4

24.6

24.6

59.7

59.3

16.5

18.3

7.7

8.7

12.1

.11.6

35.2

34.8

17.1

17.0

48

45

48

51

BIGELOW AND SCHROEDEE : SQUALOID SHARKS 119

This joint species was originally described by Gunneriis in
1766 under the name Squalus carcharias, but by the rules of
nomenclature, it must be called *microcephalus Bloch and Schnei-
der 1801, "the name Squalus carcharias having been used pre-
viously by Linnaeus 1758, for a very different shark" (Bigelow
and Schroeder 1948, p. 515). The known range of *microcephalus
extends from the White Sea, Bear Island, Spitzbergen and east
and west Greenland, southward to the North Sea (to the mouth
of the Seine as a straggler) in the east, to the Gulf of Maine
in the west.

A second North Atlantic species, rostratus Risso 1826, re-
sembles *microcephalus so closely in general appearance that
as recently as 1928 Key (1928, p. 480), following Garman (1913,
p. 241), classed it as identical with the former, while — to fur-
ther confuse the issue — recent illustrations of rostratus by
Burkhardt (1900, p. 564, fig. 3) and by Maul (1955, PI. 3, fig.
17) differ as to the length of the upper side of the caudal
peduncle relative to the length of the base of the second dorsal
fin. Both, however, like Canestrini (1864, PL 2, fig. 2) and
Ilelbing (1904, PL 9, fig. A) show the distance from the rear tip
of the second dorsal to the origin of the upper side of the caudal
as less than 70 per cent as long as the base of the second dorsal
(about as long as the base of the second dorsal in *micro-
cephalus). A more important difference is that the lower teeth are
pictured as with the cusp more nearly erect in rostratus (Fig.
15D, see also Maul 1955, PL 3, fig. 19) than they are in *micro-
cephalus (PL 3), and its denticles as rising less steeply from
the skin (cf. Maul 1955, PL 3, fig. 20, with Bigelow and Schroe-
der 1948, fig. lOlB).

Rostratus is also much the smaller member of the pair, the
greatest length we find definitely recorded for it being only
1000 mm. (Helbing 1904, p. 347) ; whereas adults of *micro-
cephalus average 8-14 feet (2.4-4.3 meters) long, with a few
growing considerably larger still (p. 116). A further difference
already remarked upon by Burkhardt (1900, pp. 561-562, 565)
is that while the luminescent organs of rostratus (p. 117) are
surrounded and thus rendered conspicuous by denticles that
differ in blunter cusps and more definitely stellate bases from
those elsewhere on the sides, this is not the case in *niicro-
cephalus, as we can verify from our own examination of a well

120 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

preserved male 1334 mm. long, taken off Massachusetts. For that
matter, we have not been able to detect anything suggestive of
luminescent organs on it, or on a 1564 mm. male from Iceland
(p. 118).

Helbing (1904), also, in his extensive treatise on the anatomy
and systematic relationships of Somniosus (as "Laemargiis'^),
has pointed out several skeletal differences between rostratus and
* micro cepJialus (as "horealis") . Finally, rostratus is ovo-
viviparous (see Helbing 1904, p. 358, for illustration of its
embrj^os at successive stages in growth), as are all the other
genera of Squaloidea for which the mode of development is
known, whereas it is still an open question whether *micro-
cephalus is ovoviviparous or oviparous (for a summary of avail-
able evidence, see Bigelow and Schroeder 1948, p. 520).

For a more detailed discussion of the differences in propor-
tional dimensions between rostratus and *microcephalus, see
Tortonese 1937-1938, p. 74.

It has recently been suggested by Maul (1955, p. 9) that
the Portuguese shark described and pictured under the name
rostratus by Capello (1870, p. 148, PI. 9, tig. 2) actually repre-
sents a third Atlantic species of Somniosus (as yet unnamed),
for it is shown and described as differing from rostratus in a
longer snout and head, and as with the position of the second
dorsal fin different in relation to the pelvics, to which we may add
a longer caudal peduncle. This appears from the following com-
parative dimensions of (A) rostratus according to Maul (1955,
measurements, p. 9, and fig. 17), and of (B) the Portuguese
specimen reported under that name, from Capello 's measure-
ments, and illustration, also measurements, contributed by Dr.
A. M. Ramalho, who kindly re-examined the specimen now in
the Bocage Museum in Lisbon.

*p^

A B

Length of head, to pectorals, in per cent of

total length 20 28-29

Snout to mouth, in per cent of length of head 25 35

Distance from rear end of base of 2nd dorsal

to upper origin of caudal, in per cent of

total length 7 11

BIGELOW AND SCHROEDER : SQUALOID SHARKS 121

Also, the origin of the second dorsal, pictured as opposite the
rear end of the bases of the pelvics by Maul, for his Madeira
specimen, is about 10 mm. posterior to the pelvics on the Capello
specimen, as we are informed by Dr. Ramalho, while excellent
photographs of its mouth, supplied by him (Fig. 15E) show its
lower teeth as somewhat more strongly oblique than they are
pictured for the typical rostratus (Fig. 15D after Canestrini,
1864, PI. 2, fig. 4; see also Maul 1955, fig. 19), i.e., resembling,
rather, those of microcephalus. Thus, present indications are that
the Capello specimen may well represent an undescribed species
more nearly related to * microcephalus than to rostratus. But
it seems to us wiser to leave the matter open than to burden
shark literature with a new name that might soon be relegated
to oblivion.

The early reports of Somniosus from the North Pacific, west
and east, were either as ^'microcephalus" or as "hrevipinna,"
on the assumption that the Pacific representative of the genus
was identical with the Atlantic ^microcephalus. And the Jap-
anese specimen, on which we based our new species pacificus
(described and pictured by Garman 1913, p. 240, PI. 15, figs.
1-3, as '"hrevipimia"), agrees with *microcephalus (p. 117) in
lacking any evident dermal structures to which a luminescent
nature might be ascribed, and in the shape of its lower teeth.
But its first dorsal stands considerably farther rearward than
on the Atlantic specimens that we have seen of *microcephalus,
or than is pictured for rostratus; the distance from the tip of
its second dorsal to the origin of the upper side of its caudal is
shorter relative to the size of the second dorsal ; the upper pos-
terior and lower anterior margins of its caudal are more convex ;
and its upper teeth are broader. It was because of these differ-
ences that Ave proposed the new species *pacificus for it (Bige-
low and Schroeder 1944, p. 35). And it seems reasonably certain
(though not yet proven) that the various reports of Som^iios us
from northeastern Asia, Alaska, and thence southward along the
Pacific Coast of America to California all were based on this
.same species.

Japanese waters also harbor a second Somniosus (Fig. 15A,
B, C, PI. 3), the *Heteroscynmus lo7igus of Tanaka 1912a (p.
102, PI. 26, figs. 102-107), more nearly resembling roslratus than
either * microcephalus or *pacificus in the shape of its lower

122 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

teeth, in the minuteness of its denticles, and in the length of
the base of its first dorsal fin relative to the length of the head.
So closely, indeed, does Tanaka's account of its type specimen,
corroborated by our examination of a female of about 1010 mm.
(in rather poor condition), received from Dr. Tokiharu Abe,
agree with the published accounts of rostratus (we have not seen
that species) that it is doubtful whether the Japanese popula-
tion differs consistently enough in any respect from the At-
lantic population to deserve a separate specific name — final
decision is a matter for the future.

Somniosus has not been reported from tropical-equatorial
waters. But the micro cephalus-pacificus division of the genus
is represented in suhantarctic latitudes by 8. antarcticus Whit-
ley 1939, a name proposed for an ''8 foot, 2 inch" (2491 mm.)
specimen that was found stranded on the beach at Macquarie
Island in latitude about 54°49'S., south of Australia (Waite
1916, p. 51, fig. 10).

The position of its first dorsal fin, as pictured by Waite, and
the shape of its caudal suggests a closer resemblance to *micro-
cephalus of the northern Atlantic than to *pacificus. But final
decision as to its specific relationship must await a more de-
tailed study of the Southern Hemisphere population.

Key to Species of Somniosus in the Northern Hemisphere

1. Lower teeth only moderately oblique (Fig. 15D) ; adults with evident
luminescent organs, at least in best known species (see p. 117)

rostratus Eisso 1826.
Mediterranean and Madeira; p.
119; also longus Tanaka 1912a,
Japan, perhaps not distinguish-
able from rostratus, see discussion,
p. 121.
Lower teeth so strongly oblique that the inner margins are almost
parallel with the trend of the jaw, forming a nearly uninterrupted
cutting edge (PI. 3) ; no evident luminescent organs 2

2. Head to pectorals 28-30 per cent of total length; lower teeth as in
Fig. 15E rostratus Capello 1870

{not of Eisso 1826), off Portugal;
see discussion, p. 120.
Head to pectorals only about 24-26 per cent of total length 3

BIGELOW AND SCHROEDER : SQUALOID SHARKS 123

3. Origin of first dorsal much nearer to tip of snout than to tip of
caudal ; interspace between first and second dorsals as long as from

snout to lpt-2nd gill openings or longer

*'microcephalus Bloch and Schneider

1801. Both sides of northern North

Atlantic, p. 117.

Origin of first dorsal almost as near to tip of caudal as to tip of

snout; interspace between first and second dorsals only about 66 per

cent as long as from tip of snout to second gill openings

*pacificus Bigelow and Schroeder
1944. Type locality Japan, prob-
ably also northeastern Asia and
Pacific Coast of North America.
from Bering Sea south to Puget
Sound, occasionally to southern
California, p. 121.

Genus ISISTIUS Gill 1864

I.<iistius Gill 1864, p. 264; type species *Scymniis hrasiliensis Quoy and
Gaimard 1824, off Brazil.

Generic characters. Dalatiinae without dorsal fin spines ; lower
teeth symmetrically triangular and smooth-edged, or only faintly
and irregularly serrate ; snout in front of mouth shorter than
from mouth to first gill openings; lip fold at corners of mouth
smooth but greatly expanded (Fig. IF; Bigelow and Schroeder
1948, fig. 98B), and supported by special cartilages that are
easily felt ; groove at corners of mouth extending rearward for
about half (44-57 per cent) of the distance toward first gill open-
ings (Bigelow and Schroeder 1948, fig. 98B) ; no preoral clefts,
but with voluminous subdermal pouches extending forward from
oral pockets (Fig. IF) ; gill openings minute, only about 4.3-4.4
per cent as long as head to origin of pectorals on w^estern North
Atlantic and Japanese specimens measured ; rear end of base
of first dorsal fin about abreast of origin of pelvics ; second
dorsal only very slightly longer at base than first dorsal (1.2
limes on specimens measured) ; interspace between first and
second dorsal fins only about 50 per cent as long as between
pelvics and lower origin of caudal; caudal fin sublunate, Avith
well-marked subterminal notch, nearly or quite as broad above
caudal axis as below, its upper margin about 1.5 times as long
as loAver anterior margin ; the caudal axis raised only very

124 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

slightly. Denticles on sides of body truncate, with concave
crown.

Maximum recorded length close to 500 mm. (Bigelow and
Schroeder 1948, p. 512).

Range. Tropical and subtropical belts of Atlantic, Pacific and
Indian Oceans.

Remarks. It has long been common knowledge that the sole
known representative of Isistius {*hrasiliensis Quoy and Gaimard
1824) is brilliantly luminescent, the entire lower surface of its
body (except for the dark collar) shining with a vivid greenish
light that presumably is produced by black dots with which
the lower surface is strewn densely; the sides, the dorsal and
caudal fins, and the basal parts of the pectorals more sparsely
so (see Bennett 1840, p. 225, for an eye-witness account of the
luminescence ; Burkhardt 1900, pp. 555, 556, 565, fig. 5 for the
distribution of luminescent dots).

Species. The various reports that have come to hand for
Isistius from one part of the ocean or another seem all to have
been based on the one species that was described first by Quoy
and Gaimard (1824, p. 198) from Brazilian waters as *Scymnus
brasiliensis. Brasiliensis has subsequently^ been taken in the Gulf
of Guinea, off Sierra Leone and Cape Verde, and north of the
Bahamas in the Atlantic ; near the Galapagos and the Hawaiian
Islands, in Japanese waters, near Fiji, west of Christmas Island,
north of New Guinea, near Lord Howe Island off New South
Wales, between Java and western Australia and near Mauritius
in the Pacific and Indian Oceans. Recent records for the mid-
Pacific are of three specimens, taken at the surface in the equa-
torial belt southward from the Hawaiian group (Lat. 2°04'S,
Long. 168°57'W; Lat. 4°47'N, Long. 161°04'W; and Dat.
2°09'N, Long. 158°14'W) where the water is 1000-3000 fathoms
(about 1830-5485 meters) deep (King and Ikehara 1956, pp.
18-19). (For a description and list of references, see Bigelow
and Schroeder 1948, pp. 509-514, figs. 98-99.)

Genus SCYMNODALATIAS Garrick 1956

Scymnodalatias Garrick 1956, p. 564; type and only known species, Scymno
don sherwoodl Aichey 1921, p. 195, PI. 39, New Zealand.
Generic characters. Dalatiinae without dorsal fin spines but

BIGELOW AND SCIIROEDEB : SQUALOID SHARKS 125

with (iistinct preoral clefts (see discussion, p. 12) ; teeth smooth-
edged, the uppers "needle-like, with asymmetrically twisted
cusps" (Garrick 1956, p. 564), the lowers with triangular cusp
much more nearly erect than in Somniosus; snout in front of
mouth only about as long as from mouth to first gill openings;
head to pectorals between 20 and 25 per cent of total length;
longest gill openings pictured as about 9 per cent as long as
head to pectorals; lips at corners of mouth not conspicuously
expanded; furrow from corners of mouth reaching rearward
only about 17-20 per cent of distance toward first gill openings ;
pre-oral clefts, described by Garrick (1956, p. 568) as "upper
labial furrows," short, but clearly shown on Garrick 's (1956,
PI. 1, figs. A, C) illustrations of type specimen; whether or not
expanded inwardly is not known. Caudal fin with distinct sub-
terminal notch ; upper margin about twice as long as lower-
anterior margin ; caudal axis moderately raised ; origin of first
dorsal fin far in advance of pelvies ; second dorsal fin a little
larger than first dorsal; interspace between first and second
dorsal fins about twice as long as between second dorsal and
caudal ; denticles on sides below first dorsal fin scale-like, over-
lapping, the free margins strongly tridentate, with median point
considerably' the longest, the outer surface with three ridges cor-
responding to the marginal points.

Size. The state of development of the claspers of a male
sherwoodi, 803 mm. long (only specimen yet seen) makes it
likely that this shark matures sexually when it is 900-1000 mm.
long.

Range. "So far known only from New Zealand.

Remarks. Archey's (1921, p. 195) original account of sher-
woodi credits it with dorsal fin spines in the form of "scarcely
discernible rudiments imbedded in the skin." Garrick (1956,
p. 556), however, on examining the type specimen was unable
to find any indication whatever of a spine in either dorsal fin,
even on dis.section, nor a.ny indication that spines might have
been removed. Seemingly what Archey interpreted as spines
actually were the "most anterior of the radial cartilages sup-
porting the dorsal fins" (Garrick 1956, p. 556). It is for this
reason that we follow Garrick in placing Scymnodalatias
among the Dalatiinae, rather than among the Squalinae, where
Archey (1921) originally located it. Actually it bridges the gap

126 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

between the two subfamilies, having a well marked preoral cleft
reaching forward from each corner of the mouth (for discussion,
see p. 12), and in having upper teeth longer in the middle third
of each side of the jaw than either toward the center of the jaw
or toward its outer corner, as is true of Scymnodon (p. 96).

Scymnodalatias resembles the Japanese *Somniosus longus
(Tanaka 1912a, p. 102) in its bodily proportions in general, as
well as in the sizes, shapes and relative positions of the dorsal,
pelvic and pectoral fins, and in the shape of its lower teeth.
But it differs sharply from all members of the genus Somniosus
in the shape of its caudal fin (see key, p. 112), and especially in
having well marked preoral clefts (see above under Generic
Characters). Other differences, seemingly generic, to which Dr.
Garrick has called our attention in a recent letter are that the
second dorsal fin is larger than the first in Scymnodalatias, that
the first dorsal stands farther rearward, and that its lower teeth
are more erect.

Species. Only one species referable to Scymnodalatias has
yet been reported, namely the Scymnodon sherwoodi of Archey
1921, so far known from a single specimen found on the beach
near Canterbury, New Zealand, in 1920. For a detailed and well
illustrated account of it, with discussion of its systematic rela-
tionships, see Garrick 1956.

Genus EUPROTOMICRUS Gill 1864

Kuprotomicrm Gill 1864, p. 264, footnote 4, type species Scynmus (Lae-

marf/u.<i) labordii Miiller and Henle 1841, p. 94, Islands of Mauritius

ri.id Bourbon, Indian Ocean; equals *Scymnus bispinatiis Quoy and

(iaimard 1824. p. 197, PI. 44, figs. 1, 2.-'

Generic characters. Dalatiinae without dorsal fin spines; the

snout blunt, its length in front of mouth about as great as from

mouth to level of third gill openings; caudal peduncle without

precaudal pits. Head to pectorals between 20 and 25 per cent

of total length ; lips at corners of mouth thick, fleshy, whether

or not with supporting cartilages is not known; no preoral

clefts, but with a voluminous pouch extending forward from

■-'In replaeiu!,' Quoy and Galmard's (1824) name *bispi)iatus with labordii,
Miiller and Henle formalized Quoy and Galmard's use of "Leiche laborde" as the
Prench vernacular name for this shark, proposed in memory of M. Theodore
r.aborde.

BIGELOW AND SCHROEDER : SQUALOID SHARKS 127

each oral pocket; groove from corner of mouth reaching rear-
ward for 50-66 per cent of the distance toward first gill open-
ings; gill openings minute (Fig. 16F). First dorsal fin con-
siderably posterior to mid-length of trunk, but its base well in
advance of origin of pelvics; second dorsal about 2.5 times as
long, at base, as first dorsal ; interspace between first and second
dorsals a little longer than between second dorsal and caudal;
caudal fin without subterminal notch, its upper margin about 1.2
times as long as lower-anterior margin; caudal axis moderately
raised and continuing to extreme rear margin of the fin ; the fin
wider below the axis than above; dermal denticles described as
low, quadrate, wdth concave crowns and of two sizes (Hubbs and
McHugh 1951, p. 167), the smaller much the more numerous,
the larger with stellate bases, and with additional pits surround-
ing the axial concavity."* Teeth smooth edged ; the uppers
slender, sj^mmetrical, slightly recurved ; the lowers with cusp
directed obliquely outward, the successive inner edges at an
angle of about 45° with the jaw.

Luminescence. The single known species of Eiiprotamicrus is
one of the few sharks that are known to be luminescent, glow-
ing, as described by Dickens (1956), with a greenish light that
is given off from its lower surface by "manj^ thousands of small
light organs," according to Marshall (1956, p. 73).

Size. Lengths of 233 mm. reported by Hubbs and McHugh
(1951, p. 170) and of "8-9 inches" by Giinther (1870, p. 428)
for the only known species of the genus probably approximate
the maximum to which it ordinarily grows, for Dumeril (1865.
}). 457) described the claspers of a male of 205-210 mm. as having
a strong spine on the outerside, i.e., as close to maturity. Simi-
larly, the gravid female, 234 mm. long,^^ recently reported by
Dickens (1956) and by Marshall (1956) gave premature birth
to six young when brought on board ship. Thus Euprotomicrus
falls in the same size-group with Squaliolus (p. 129) and -with
the smaller of the species of Etmoptcrus (p. 48).

Species. All the various reports for Euprotomicrus (see Gar-
man 1913, p. 235 for synonyms under hispinatus Quoy and Gai-

2S We have detected the smaller denticles alone on the specimen we have seen.
;ind only a few of these, here and there. Seemingly its skin has been largel.v
denuded of its armature.

29 Information contributed by Mr. N. B. Marshall.

! I'b BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

mard 1824 and E. hyalinus Eigenmann 1891) seem certainly to
have referred to the type species Mspinatus Quoy and Gaimard
1824, of which we have been fortunate enough to study a female,
161 mm. long, from the North Pacific, west of Johnston Island.
For a description of it, with measurements, see King and Ike-
hara 1956, p. 17. For a description of a somewhat larger female
(233 mm. long) taken in 1948 some 500 miles off California,
with excellent illustrations, and wdth discussion of the relation-
ships of the genus, see Hubbs and McHugh 1951. The most
recent report is that of a gravid female 234 mm. long, taken
from the SS ''Kent" in the southern Indian Ocean west of
Australia, Lat. 23°10'S., Long. 101°58'E. (Dickens 1956; Mar-
shall 1956).

E. hispinatus is known from the Indian Ocean, near Mauritius
and Bourbon as well as west of Australia (see above), from
New Zealand waters, from the northern Pacific localities men-
tioned above, and from between Honolulu and San Francisco
(Eigenmann 1891, p. 35, as E. hyalinus). Our earlier reference
of the genus to the Philippines (Bigelow and Schroeder 1948, p.
500) was based on the misconception that the generic name
Sqnnliolns Smith and Radcliffe 1912 is an equivalent.

Evidently this is a pelagic species; the specimen we have
seen (see above), the one described by Hubbs and McHugh and
the one reported by Capt. Dickens (1956) having all been picked
up at the surface at night, where the water is more than 2000
fathoms (about 3660 meters) deep.

Genus SqUALIOLUS Smith and Radcliffe 1912

Squaliohis Smith and Radcliffe, 1912, p. 683. Type species *S. laticaudua
Smith and Radcliffe 1912, p. 685, PI. "0, Philippines.
Generic characters. Dalatiinae with a short spine in the first
dorsal fin, either exposed at the tip (Fig. 16A, D) or wholly en-
closed in the skin, but without a spine in the second dorsal fin ;
snout in front of mouth nearly or quite as long as from mouth
to origin of pectorals ; head, to pectorals, al)out 33 per cent of
total length ; caudal peduncle without precaudal pits ; lips at
corners of mouth thick and fleshy, as in Euprotomicrus (p.
126) ; no preoral clefts, but w4th a voluminous pouch extending
forward from each oral pocket (Fig. 16C) ; furrows from corners

BIGELOW AND SCHROEDER : SQUALOID SHARKS 129

of mouth reaching rearward 50-70 per cent of the distance
toward first gill openings (Fig. 16 A, B). Base of first dorsal
spine considerably posterior to a perpendicular at axil of pec-
torals; second dorsal fin at least twice as long at base as first
dorsal, its rear tip reaching to a point about midway between
rear end of base of second dorsal and origin of upper side of
caudal. Caudal fin sublimate, with bluntly pointed tips and
without subterminal notch ; upper margin only slightly longer
than lower anterior margin ; the maximum width of the fin
nearly as great above the axis as below (Fig. 16 A) ; caudal axis
not raised, except that the extreme tip curves upward at the
margin of the fin (Fig. 16A). Teeth smooth edged, uppers
slender, sj^mmetrical, somewhat recurved, lowers with cusps
directed obliquelj' outward, the inner edges weakly concave in
outline, making an angle of about 33° with the jaw (Fig. 16C) ;
the general contour of the tooth deeply notched on the outer side
marking the transition from cusp to base. Dermal denticles
rather closely spaced and so prominent that the skin has a
pebbled look (PI. 4), truncate, with concave crowns (Fig. 15H),
on four-radiate bases, and varying in size from denticle to
denticle.

Size. The type species, laticaudus, matures when only about
150 mm. long, to judge from the stage in development of the
claspers of the type specimen (Smith and Radelitfe 1912). Thus
Sqiialiolus includes the smallest shark for which the size at
maturity is known, though it is closely rivalled by Eicprotomicrus
(p. 127), b}^ the smaller of the species of Etmopterus (p. 48) ;
also by the smallest of the members of the family Scyliorhinidae
among the galeoid sharks. Sarmenti, the second known species
of Squaliolus, grows somewhat larger, the two specimens yet
seen having measured 224 mm. and 226 mm. in length (both
were females).

Depth range. The two specimens of the type species {lati-
caudus) that have yet been seen were taken in Philippine waters
in a beam trawl dragging at about 310 meters (170 fathoms,
Smith and Radcliffe 1912, p. 685) ; they may, of course, have
been picked up by the trawl either on its way down or on its
way up again (p. 5). The case of the second known species
is puzzling, for while the original specimen was caught near
Madeira on a long line, presumably fishing at least as deep as

lao

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

P
O

l-H

BIGELOW AND SCHROEDER : SQUALOID SHARKS 131

900 meters or so (Noronha 1926), the second known specimen
was caught in very shoal water, near Arcachon, France, among
eel grass (Zostera). (For details see Sigalas 1939-40, p. 70.)

Remarks. In our earlier synopsis of the dalatiine sharks (Big-
elow and Schroeder 1948, p. 500, footnote 2) we classed Squalio-
lus as a synonym of Euprotomicrns, as had Garman (1913, p.
234) and Fowler (1941, p. 264). Recent examination, however,
of the type specimen of laticaudus (in the U. S. National Mu-
seum), and of the female pictured on Figure 16 A has convinced
us that Squaliolus is separable generically from Euprotomicrns
from which it is set apart not only by the presence of a spine in
the first dorsal fin (it differs in this from all other Dalatiinae for
that matter), but also by the position, much farther forward,
of its first dorsal fin, by its much longer and more pointed snout,
and by its conspicuously longer head.

In the type species each nostril is prolonged at the outer end
by a conspicuous groove directed forward, and at the inner end
by a similar groove directed rearward (Fig. 16B). But we
hesitate to include this feature among the generic characters,
because the condition in this respect is not known for the second
known representative of the genus {S. sarmenti, see below).

Species. Two species referable to Squaliolus are known, *8.
laticaudus Smith and Radcliffe 1912, (from the Philippines),
type of the genus, and S. sarmenti Noronha 1926, which was
originally described from Madeira, and which has recently been
reported from Arcachon, France, by Belloc (1937) and by
Sigalas (1939-40). The chief difference between these two is that
while the tip of the first dorsal spine is exposed in ^laticaudus
(Fig. 16 A, D), it is described by Noronha as entirely enclosed in
the skin in sarmenti. Other differences mentioned by Noronha

Fig. 16. A, SqxMliolus laticaudus, female, 143 mm. long, Philippines,
U. 8. Nat. Mus., No. 70259. B, ventral view of anterior part of head of
same. C, right hand comer of mouth of same, the preoral pouch indicated
by the broken curve, x about 4. D, first dorsal fin of same, to show spine, x
about 3. E, Heteroscymnoides marleyi, after Fowler, slightly emended from
comparison with the type specimen, a female, 126 mm. long, from the
Durban Coast, southern Africa (see p. 133). F, Euprotomicrns hispinatus,
female, 161 mm. long, from central Pacific west of Johnson I., U. S. Nat.
Mus. No. 164176.

132 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

are that the first dorsal fin stands a little farther forward in
sarmenti, in which the distance from snout to first dorsal origin
is about 42 per cent as long as the trunk (about 48 per cent in
Haticaudus) , and that the pectorals, when laid back, reach only
about as far as the concealed first dorsal spine in sarmenti, but
overlap the base of the fin in Haticaudus. It may also prove
that S. sarmenti lacks the grooves extending from the inner and
outer ends of the nostrils that characterize *S. laticaudus, for
they are not indicated on the illustrations of sarmenti (Noronha
1926, PI. 35). But the latter are not clear in this respect.

Genus HeTEROSCYMNOIDES Fowler 1934

Heteroscymnnides Fowler 1934, p. 239, type species *H. marleyi Fowler
1934, p. 240, Dur1)an coast, southeastern Africa.

Generic characters. Dalatiinae without fin spines; snout in
front of mouth about as long as from mouth to third gill open-
ings; head (to pectorals) more than Ys of trunk (to origin of
caudal) ; gill openings minute, not more than 8-4 per cent as long
as head to origin of pectorals ; lip at corners of mouth neither
conspicuously thick and fleshy nor expanded ; furrow at corners
of mouth reaching rearward about 40 per cent of the distance
toward first gill openings ; no preoral clefts ; whether or not with
preoral pouches is not known ; origin of first dorsal fin close
behind a perpendicular at origin of pectorals ; second dorsal fin
about 1.2 times as long (at base) as first dorsal, its origin
over bases of pelvics ; distance from rear base of second dorsal to
origin of upper side of caudal fin about 50 per cent as long as
interspace between first and second dorsals; caudal (in excellent
condition on the one known specimen) without subterminal
notch, about 1.7 times as wide below caudal axis as above; upper
margin about 1.3 times as long as lower anterior margin; caudal
axis not appreciably raised. Dermal denticles on sides of
body (Fig. 15G) triangular spur-like, with three prominent
ridges, raised onlj^ slightly, on quadrate bases, and so closely
spaced that the skin is almost wholly concealed. Upper teeth
erect, lowers directed somewhat obliquely outward (Fowler
1941, p. 273).

The size at maturity is not known, the only specimen yet seen
being a newly born female, 126 mm. long, with healed umbilical

BIQELOW AND SCHROEDER : SQUALOID SHARKS 133

scar. Presumably this was from shoal water, but the depth of
capture is not known.

Remarks. The features that set Heteroscymnoides the most
obviously apart from related genera are its long snout and head,
minute gill openings, shape of caudal fin, and location of the
first dorsal fin.

Species. One species only (the type) is known, *H. marleyi
Fowler 1934. For a detailed account, with illustration showing
its general appearance, see Fowler's (1934, p. 239; 1941, p.
273) original description of the only specimen that has yet been
seen. Having had an opportunity to examine the latter through
the kindness of Dr. James Bohlke of the Academy of Natural
Sciences, Philadelphia, we include the following table of its pro-
portional dimensions as per cent of total length, with number of
teeth.

&novt, length in front of : eye

11.9

moutli

17.1

Eye: horizontal diameter

4.8

Mouth: breadth

8.0

Nostrils: distance between inner ends

4.0

Gill openings: length of longest

1.0

First dorsal fin: vertical height

3.2

length of base

6.4

Second dorsal fin : vertical height

2.8

length of base

8.2

Caudal fin : upper margin

18.3

lower anterior margin

13.9

Pectoral fin: outer margin

10.7

Distance from snout to: first dorsal

34.1

second dorsal

65.8

upper caudal

81.7

pectorals

30.1

pelvies

57.5

Interspace betu-ecn: first and second dorsals

22.2

second dorsal and caudal

10.0

Distance from origin to origin of : pectorals and pelvies

27.4

pelvies and

caudal

21.4

Teeth:

6-1-6

11-1-11

134 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Subfamily ECHINORHININAE

Squalidae without fin spines; the teeth with several cusps,
the median cusp much the largest; upper teeth similar to lowers;
caudal axis raised, the fin wider below the axis than above,
its tip pointed, without subterminal notch ; short preoral clefts,
but not expanded inwardly, nor is the pocket at each corner of
the mouth extended rearward, furrow-like, as it is in all other
known representatives of the family Squalidae ; dermal denticles
either simple, with central cusp, or compound, with two or more
cusps, and varying correspondingly in size on any given in-
dividual ; lateral line, along sides of body roughened with series
of minute thornlets (see Remarks, below). Recorded up to 8-9
feet long.

Echinorhiniis Blainville 1816 is the only known genus.

Genus ECHINORHINUS Blainville 1816

Echmorhinns Blainville 181(i, p. 121 ; type species E. spinosus, no author
quoted but doulitless the *Squalus hrunis of Bonnaterre 1788, p. 11.
intended.
For generic synonyms, see Bigelow and Schroeder 1948, p. 526.

Generic characters. Those of the subfamily.

Depth range. "In all warm seas from 10-100 fathoms" (18-
183 meters), Smith 1949, p. 56.

Remarks. The combination of characters given above under
the subfamily, especially the buckler-like denticles, often with
2 or more cusps, and the peculiar armature of the lateral line,
would place Echinorhiniis at a glance, should one come to hand in
any part of the world where it is not well known.

Accounts of the lateral line in Erhinorhinus have been con-
flicting. Rey (1928, p. 488) who had seen only a dried skin,
described it as in the form of a deep furrow enclosed between
two close-set rows of slender tentacular structures or dermal
papillae, though with the reservation that he could not form an
exact idea of it. This is in line with McCoy's (1887, PI. 144,
fig. IG, reproduced by Whitley 1940, fig. 169) illustration of it
as flanked by two sparse rows of flexible papillae. And it was
from these sources that our earlier account of it (Bigelow and
Schroeder 1948, p. 527) was drawn. McCoy (1887, p. 165), how-
ever, described it for his Australian specimen as "roughened

BIQELOW AND SCHROEDER : SQUALOID SHARKS 135

with rows of minute, slender hooked spines" as far forward as
the thii'd gill openings. And it is similarly armed with minute,
curved, thorn-like spines on an excellent specimen from the
Mauritanian Coast that we have received through the kindness
of Dr. Ramalho, the only suggestion of papillae or the like being
that the course of the channel is marked by numerous shreds of
skin and flesh, seemingly the result of wear. In its present state
the canal is open along most of its length on both sides of the
body. But as seen in cross-section its lower lip overlaps the
upper. And since it is bridged over here and there for short
stretches by the tissue-paper-thin skin, we assume that normally
the canal is closed, at least as far rearward as the pelvic fins,
if not farther still. A touch is enough to show that the spinelets
are denticular structures, not fleshy. And they are in plain view
(under a lens) wherever the canal is open. But we have not
been able to determine (because of the condition of the speci-
men) whether they are exposed at the tips along the short
stretches where the canal is roofed over, or whether they are
entirely enclosed, as is suggested both by surface views and by
cross sections of the canal.

Species. The following named species fall in Echinorhinus:
^Squalus hrucus (Bonnaterre) 1788 (also reported repeatedly
as spinosus Blainville 1825), the type of the genus, well known
in the eastern North Atlantic; Echinorhinus ohesus Smith 1849,
PI 1, South Africa ; E. cookei Pietschmann 1928, p. 297 ; 1930,
p. 3, Fig. 1, Hawaiian Islands; and E. (Ruhusqualus) niccoyi
Whitley 1931, p. 311, 1940, p. 151, Victoria, Australia. But
Fowler (1941, p. 278) who examined a specimen of cookei in
Honolulu could not find "that Echinorhinus cookei is other than
a variant of *hrucus", nor can we find anything in the accounts
of ohesus or of mccoyi to suggest that they differ significantly
from *brucus, as represented bj^ a specimen of the latter from
the eastern Atlantic in the British Museum (Bigelow and
Schroeder 1948, Fig. 102), and by a male 1600 mm. long, from
the coast of Mauritania, recently received through the kindness
of Dr. A. M. Ramalho. On this basis, the recorded range of
*hrucus includes the eastern North Atlantic from tropical West
Africa to Ireland and the North Sea (including the Mediter-
ranean) ; the coast of Massachusetts in the western North At-

136 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

lantic (one specimen found washed ashore at Provincetown,
Massachusetts, in 1878); South Africa; Argentina; California;
the Hawaiian, Japanese, and Australo-New Zealand regions, and
Arabian waters. (For further details and for references, see
Bigelow and Schroeder 1948, pp. 529-532.)

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PLATES

Plate 1. Oxynotus centnna. Above, lower surface of anterior part of head
of female, 510 mm. long, coast of Portugal, x about 1.5. Below, roof of
mouth of male, 590 mm. long, Portugal, viewed from below, to show
arrangement of upper teeth, x about 6, both specimens Mus. Comp, Zool.
No. 39576.

PLATE 1

Plate 2. Etmopierus bulUsi, new species. Female, 196 mm. long, off
northeast coast of Florida, Pelican sta. 42 (see p. 50). Above, side view
of mid-sector of trunk, x about 3, to show the linear arrangement of the
dermal denticles. Beloiv, ventral view of same, at pectoral fins, to show the
random arrangement of the denticles on the lower surface.

PLx^TE 2

Plate 3. Above, Somniosus longus, female 1010 mm. Mus. Comp. Zool.
No. 39650, upper and lower teeth, left hand side of mouth, x about 3.3.

Middle and helow, Somniosus microcephalus, upper and lower teeth, along
central part of mouth, from jaws of a Gulf of Maine specimen about 11 feet
long, Mus. Comp. Zool. No. 36119, x about 3.

PLATE 3

Plate 4. Above, Squaliolus laticaudus, same speeiiiien as in Figure 16A,
photograph of mid-portion of trunk, to show papillate appearance of skin,
X about 2.8.

Below, Euprotomicrus hispinatus, same specimen as in Figure 16F, corre-
sponding photograph to show the smoothness of the skin as contrasted with
Squaliolus (above), x about 3.

PLATE 4

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 117, No. 2

CHECK LIST OF THE REPTILES AND AMPHIBIANS

OF EAST AFRICA (UGANDA; KENYA;

TANGANYIKA; ZANZIBAR).

By Arthur Loveridge

CAMBRIDGE, MASS., U. S. A.

printed for the museum
August, 1957

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 18G3 — The current volume is Vol. 117.

Breviora (octavo) 1952 — No. 80 is current.

Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55.

Johnsonia (quarto) 1941 — A publication of the Department of Mollusks.
Vol. 3, no. 35 is current.

Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 2, no. 21 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 numbers
which may be purchased separately. Prices and lists may be obtained on
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Of the Peters "Check List of Birds of the World," volumes 1-3 are out
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published under Museum auspices.

Bulletin of the Museiun of Comparative Zoology

AT HARVARD COLLEGE

Vol. 117, No. 2

CHECK LIST OF THE REPTILES AND AMPHIBIANS

OF EAST AFRICA (UGANDA; KENYA;

TANGANYIKA; ZANZIBAR).

By Arthur Loveridqe

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

August. 1957

No. 2. — Check List of the Reptiles and Amphibians of East
Africa (Uganda; Kenya; Tanganyika; Zanzibar)

By Arthur Loveridge

CONTENTS

Page

lutroduction 153

Acknowledgements 162

List of the Reptiles (Turtles; Crocodiles; Lizards; Snakes) 163

List of Amphibians (Caeeilians; Toads; Frogs) 305

List of species still unrepresented in the Museum of

Comparative Zoology 358

Bibliography 360

Index to all scientific names mentioned i

INTRODUCTION

"A check list is the lowest class of literature." Though more
profound observations have long since faded from memory, the
preceding remark — made to me by a former colleague at the
National Museum of Wales — has, strangely enough, never been
forgotten. Possibly I was startled by G. R. Brook's dictum as the
idea of a cheek list being considered as literature had never oc-
curred to me. Nevertheless, however classified, a check list may
serve a useful purpose in proportion to the time and care devoted
to making it as free from error as may be humanly possible.

The present list owes its origin to a simple compilation of names
extracted from such literature as was available to me in 1914,
augmented by records of any specimens that had come my way. It
was intended solely as a working basis for my personal use when
I should reach East Africa. During the succeeding decade a
number of additions resulted from my own field work. Upon my
return from East Africa in 1923, I was urged by the then curator
of reptiles at the British Museum, Dr. Joan B. Procter, to make
it available to other workers.

Somewhat reluctantly I did so the following year, assuming I
had seen the last of East Africa since I was proceeding to the
United States. That check list, of which I never saw proof as it

154

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

was published in Kenya, certainly merited Brook's description.

At that time, however, since it was the first list to cover the area,

it was probably better than nothing!

The area covered w-as precisely the same as in the present list,

viz :

Kenya Colony and Protectorate . . 224,960 square miles

Tanganyika Territory 360,000

Uganda Protectorate 93,981

Zanzibar Protectorate' 1,020

Giving a total area of 679,961 square miles

It is, therefore, but little more than a quarter the size of the
continental United States," whose herpetofauna has been the sub-
ject of intensive study during the past century. East Africa,
whose interior was largely a terra incognita sixty years ago,
though only about a tenth of the area covered by the North
American check list, straddles the equator. Consequently, though
solely on an area basis, it enjoys a relatively richer herpetofauna,
as shown bv the following figures.

Group

Total number of forms listed as
occurring in

East Africa
in 11124 3

Kast Africa
in 1957

North

.Vnierica
in 111.").'', I

Turtles and Tortoises

13

15

81

Crocodilians

o

3

2

Lizards and Chameleons

155

179

148

Snakes

139

161

275

Amphibians

128

169

262

TOTALS

437

527

768

1 Including Pemba Island with an area of approximately 380 square miles.

2 3,022,387 square miles which, added to that of Canada (3,695,189 square
miles) gives a total of 6.717,576 for North America — the area covered bv tht
1953 check list.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

155

As so many species are common to more than one territorial
area, a regional breakdown of the new figures naturally results
in some duplication. So far as Zanzibar Island is concerned, the
situation remains obscured by the earlier use of the word Zanzibar
(== Negro coast) in a broader sense that covered the mainland
littoral of Tanganyika Territory only forty miles away. If only
some resident naturalist would conduct a thorough survey of the
island's herpetofauna, the matter might be cleared up once and
for all. Though the figures for the three offshore islands corre-
spond fairly closely to those of Moreau and Pakenham (1941:
109), actually a number of names have been dropped on account
of synonymy or for other reasons, while others have been substi-
tuted or added.

Grouii

Uganda
I'rotec-
torate

Kenya
Colony

Tan-
ganyika
Territory

Peniba
Island

Zauziliar

Island

Matia
Island

Turtles and Tortoises

7

10

12

1

5

1

Croeodili.'Uis

o

i

o

0

1

0

Lizards and
Chameleons

41

92

123

10

18

10

Snakes

69

89

113

9

33

12

Caecilians

0

3

7

0

0

0

Toads and Frogs

44

Go

114

6

21

9

TOTALS

1(53

260

371

26

78

32

Apart from the inflation of the figures for Zanzibar's herpeto-
fauna for the reasons given above, offset to some extent by the fact
that the reptiles of its dependent island Pemba are little known,
it is clear that there is a definite relation between speciation and
area.

3 The Amphibia are taken from a l!t30 list published in the Proceedings of
the Zoological Society of London, pp. 7-32.

* As no figures are given in the Check List of North American Amphibians
and Reptiles, by K. P. Schmidt (ed. 0. University of Chicago Press), these are
my own count in which are included the omitted forms in the addenda (1954,
Copeia, pp. 304-30G), but exclusive of recently introduced lizards (8 kinds)
and frogs (3 forms).

156

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

At the time of the 1924 check list, when I arrived at the Mu-
seum of Comparative Zoology, that institution had only 161 of
the 527 species and races now recognized. Today all but 33 are
represented.

Number of listed forms represented in the Museum of Comparative Zoology

Group

Present
in 1923

Additions
since 1924

Still
lacking

Total

Turtles and Tortoises

11

4

0

15

Crocodilians

3

0

0

3

Lizards and Chameleons

49

121

9

179

Snakes

61

95

; )

161

Caecilians

0

9

u

9

Toads and Frogs

37

104

19

160

TOTALS

161

333

33

527

Of this total of 527, 1 personally collected 413 in the field, while
most of the remaining additions were obtained in exchange for
duplicate material secured on the field trips I was privileged to
carry out. The majority resulted from the itineraries being care-
fully planned to embrace not only type localities of our desider-
ata, but also type localities of many species or races that were
inadequately described, or known only from one or two indi-
viduals. Topotypic series of these creatures resulted in 180 (73
+ 107) alleged forms of reptiles and amphibians being relegated
to the synonymy. Not always justifiably, though usually so.
Nevertheless, anyone who proposes describing additions to the
herpetofauna of East Africa, would be well advised to scrutinize
with care the synonyms attributed to the species most nearly
related to the allegedly new form.

It is important to note the distinction between the names listed
in the generic synonymies, and those of species or races. Follow-
ing the citations for the 135 genera (110 of reptiles; 25 of
amphibians) recognized in the present list, are 527 additional

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 157

citations. While the majority are straight synonyms, with a
sprinkling of nomina niida, others represent well-defined groups
regarded by me as of subgenerie status. This is, of course, a mat-
ter of opinion, and to some of them a few of my fellow workers
may wish to accord generic rank. Intensive study of any small
group is apt to result in observed differences assuming an undue
importance, at least temporarily. To adjust one's perspective it
is advisable to evaluate the situation on a family basis.

In the 1924 check list no synonymies were supplied, so I trust
their inclusion in the present one may serve a useful purpose. Is
it too much to hope they may act as a deterrent to reckless de-
scribers like the late Ernst Ahl, of whose 95 "species" only 28 are
considered valid? It is always possible that additional material
may demonstrate that an occasional "synonym" represents a
recognizable form, but "more material" is just as likely to have
the opposite result. That 527 recognized forms should have 830
synonyms does not quite reflect the position, for more than half
(274) have no synonyms whatever; on the other hand, several
widely distributed species have a dozen or more each. It is but
fair to say, however, that of these 1349^ more or less useless names
burdening the literature, not one was proposed by an African, but
then neither was a single valid one !

This should not be construed as a reflection, for it is little more
than a century since any one of the many indigenous tribes of the
region had a written language. Not until 1848, 1 am informed, was
the first of almost a hundred tongues in this area reduced to
writing by a white immigrant. After all, following 300 years of
civilizing contacts, it is doubtful whether an indigenous Indian
described a single one of the 768 reptiles and amphibians that
figure in the latest North American check list.

Genera. Chiefly for the benefit of herpetologists in Africa,
working without benefit of library facilities, I have listed under
each genus all of its synonyms known to me. After this check
list was typed, I was given (through the courtesy of Dr. A. S.
Romer) the opportunity to scan the extensive synonymies in an
advance copy of his "Osteology of the Reptiles" (pp. xxi -\- 772 :
Chicago: 1956). This has enabled me to add between 20 and 30
synonyms of world-wide genera to the already lengthy lists. Gen-

5 527 generic -|- 822 species or races.

158 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

erally speaking, I have avoided inclusion of variations due to
misprints or misspellings, together with some emendations (cf.
footnote to Chamaeleo on p. 36. A few names that appear in
the "Osteology" are omitted after due consideration. Unques-
tionably, further additions will have to be made in the case of
wide-ranging genera with species in Europe or Asia.

Many tedious hours have been spent in attempting to ascertain
the correct genotypes for each name. Where no type species had
been mentioned I have occasionally selected one with the words
"by present designation." Such action should be regarded as
tentative, for in some instances further research may reveal that
a type has already been proposed in some publication I have
overlooked. Had I but realized at the outset how much time this
uncongenial task would take I might well have shirked it, invok-
ing the elder Agassiz's wise counsel to "Study specimens, not
l)Ooks." Sound advice which he apparently overlooked when, in
compiling his "Nomenclature Zoologicus" (1842-1847), he pro-
posed innumerable emendations in the spelling of generic names.

However, the chief offenders were Fitzinger and Gray who, ap-
parently vying with each other in creating monotypic genera,
have each proposed about 90 names — in so far as this list is con-
cerned. Of these only 14 and 11 respectively are here regarded
as full genera. Their percentages of validity (15% and 12%)
compare very unfavorably with the more discriminating work of
Giinther (10 out of 31 valid) and of Peters (8 out of 24), each
of whom achieved approximately 33 per cent.

Fitzinger, taking some work like the "Erpetologie Generale, "
gave Dumeril and Bibron as the authors of his type species. As
they in turn attributed the authorship to Linnaeus or some other
early worker, I have cited the original author of the name with a
view to avoiding much unnecessary printing. For example, in the
case of Psammorrhoa Fitzinger (1843), instead of saying : "Type
by original designation: Agama aculcata Dumeril & Bibron =
Agama aculeata Merrem," I have cited Merrem only, omitting
Dumeril & Bibron. I ask forgiveness of those taxonomically-
minded colleagues who may be shocked by such action.

In the course of compiling these generic synonymies, I came
across several names that had priority over those currently in use.
RuliuR-s setting aside these older names should be obtained from

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 159

the International Commission on Zoological Nomenclature with-
out delay. They are :

Thermophilus Fitzinger (1843), not Thermophila Hiibner
(1816), antedates Ichnotropis Peters (1854) which has been
in use as a genus of the LACE RTI DAE for over a century.
Ibiba Gray (1825) antedates Boiga Fitzinger (part:1826), both

having the same type : Coluber irregularis Merrem.
Cerastes Laurenti (1768), long associated with certain vipers, is
quite unsuitable as a generic name for the docile opisthoglyphs
currently known as Psammophylax Fitzinger (1843), both gen-
era having the same type : Coluber rhombeatus Linnaeus.
Philodcndros Fitzinger (1843), a most inappropriate name for
the terrestrial snakes which have been known as Dromophis
Peters (1869) for almost a century. Both genera have the same
type : Dendrophis praeornata Schlegel.
Chloroechis Bonaparte (1849) antedates Atheris Cope (1862), a
name that has been in use for almost a century. Both genera
have the same type : Viper a chloroechis Schlegel.
At this point I might also mention Cobra lachesis Laurenti
(1768). This is an earlier name for Bitis a. arietans Merrem
(1820), the common Puff Adder which ranges from the Cape to
Morocco. Some years ago Cobra was rejected as generically ap-
plicable. It is regrettable that lacJiesis was not dealt Avith at the
same time, for reasons I have given elsewhere and have repeated
briefly in a footnote to arietans.

Eminophis Werner (1924, Sitzb. Akad. Wiss. Wien, 133, Abt.
1, p. 55), whose type (E. lineolata Werner) was said to have
come from East Africa, was actually based on an Indian species
— TracJiiscJium fuscum (Blyth) — according to Malcolm A.
Smith (1928, Ann. Mag. Nat. Hist., (10) 1, p. 496).

Species and Races. The 1924 check list was largely binomial,
only 7 "varieties" (of Boulengerian nomenclature) and 28 sub-
species being cited. Of the 527 reptiles and amphibians figuring
in this new list no fewer than 259 are expressed as trinomials —
a change reflecting a better understanding of relationships and
ranges. Several names appearing in the earlier list, or subse-
quently recorded from East Africa, have been omitted after an
examination of the material on which they w^re based revealed
misidentifications — usually of a subspecific nature. It is to be

160 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

hoped that they will not be reinstated by enthusiastic amateurs
without due cause. Various species or races, rather too numerous
to list, are now synonymized for the first time.

It Avill be observed that citations are given in a carefully
standardized format. Precise date of publication is sometimes a
problem. For example, it maj^ be noted that the dates I give for
many of the 34 species (common to East and South Africa)
described by Sir Andrew Smith will be found to differ from
those given by Boulenger in his catalogues. My dates were ob-
tained from a paper by F. II. Waterhouse (1880, Proc. Zool. Soc.
London, pp. 489-491), "On the Dates of Publication of the parts
of Sir Andrew Smith's 'Illustrations of the Zoology of South
Africa.' " The titles of some journals bear the name of the city
of publication, e.g., Bull. Soc. Philom. Paris. Where omitted,
and its absence may create difficulties, it has been added in
parentheses, e.g., Bull. Mus. llist. Nat. (Paris). Where a figure
carries no number or letter, to show that it has not been in-
advertently omitted I have expressed it thus : Fig. — .

It seems scarcely necessary to point out that capitals for
specific or subspecific names are no longer in general use. Like
hyphenated names they appear that way in the synonymy only
when that was the way they were originally printed ; for example,
a specific name like adolfifriderici may have appeared as Adolfi-
frider-ici in the first instance.

The description of Gecko tubercidosus Daudin (1802) seems
to agree with the characters of the Common House-Gecko of East
Africa long known as Hemidactylus mabouia (Moreau de Jonnes :
1818). However, the latter name having been in continuous use
for 150 years, it is to be hoped a ruling to set tubercidosus aside
will be secured from the International Commission on Zoological
Nomenclature.

The only outstanding change meriting mention here is the use
of Boaedon fuliginosiis fidiginosus (Boie:1827) for the Common
House-Snake, long known in East and South Africa as Boaedon
lineatus lineatus Dumeril & Bibron (1854). As the eastern rep-
tile appears indistinguishable from the western, this vexatious
change seems unavoidable.

Many extraterritorial species, even genotypes that are alien to
Africa, are mentioned in footnotes or elsewhere. All are recorded

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 161

in the comprehensive Index which is the key to a mine of infor-
mation that might he otherwise overlooked.

Ranges. In general, ranges are presented in a standardized
format, the included countries — commencing with Spanish
Morocco — being taken clockwise around the continent. Circum-
stances occasionally make deviations advisable as, for example
where forest forms of western Kenya Colony extend due west
to Gambia. Recent changes in political status such as Sudan
(for the former Anglo-Egyptian Sudan), still differentiated by
its spelling from the French Soudan, and Somalia (for Italian
Somaliland) have been followed. Eritrea, however, though now
merged with Ethiopia is faunistieally so different that it ap-
peared advisable to retain it as a separate entity. Other prov-
inces that have been similarly treated are the huge units compris-
ing the Union of South Africa, and sometimes it has seemed best
to be specific with areas like the Gabon, instead of blanketing
them in the vast region known as French Equatorial Africa. The
territories comprising French West Africa also are at times listed
individually.

When an animal occurs in any one of the four countries
covered by this check list, that country is mentioned specifically.
Though Mafia Island, lying south of Zanzibar, forms part of
Tanganyika Territory, zoogeographically it is simpler to list it
along with Pemba and Zanzibar Islands, from whose sultanate it
was once administered. The v\-ord "coastal" is employed to de-
note the lowland zone extending from the Indian Ocean to the
continental plateau. In width it varies considerably for a dozen
degrees south of the equator. Its characteristic herpetofauna
reaches inland as far as Lake Nyasa and even beyond.

It need hardly be pointed out that the ranges furnished are
only as of 195G, and subject to extension as our knowledge in-
creases. Caution should be exercised in extending them to include
improbable records, for the distribution of many species is con-
ditioned by well-defined habitat preferences. The nine principal
faunal zones have been discussed elsewhere (Loveridge, 1937,
Bull. Mus. Comp. Zool., 79, pp. 481-541), and the majority of
species in this check list assigned to one or more of the zones.

Our understanding of Hyperolius is so far from complete that
to avoid misunderstandings as to the ranges of these sedge- or

162 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

tree-frogs I have departed from the format employed elsewhere
by indicating' with an asterisk (*) precisely those countries from
which material of the particular species or race is present in the
collections of the Museum of Comparative Zoology. As a further
safeguard, countries included on the basis of unique records are
followed, in parentheses, by the name of the responsible author-
ity and the year in which he published the record. This pro-
cedure has also been used for members of the RANIDAE.

Footnotes. Primarily for the benefit of isolated workers in
Africa, explanatory annotations have been given in the form
of footnotes. These provide references to, or the reasons for, rela-
tively recent taxonomic changes ; others are intended to elucidate
ambiguities or to account for seeming omissions, especially where
due to erroneous records. A few refer to unsolved problems and
indicate profitable lines of investigation that need to be under-
taken.

ACKNOWLEDGEMENTS

The policy of submitting a check list, prior to publication, to
the scrutiny of one's colleagues is an admirable one. Unfortu-
nately, most of those interested in African herpetology are over-
seas, besides which the time factor militates against putting the
idea into practice in this instance. However, help has been
received from various correspondents and colleagues, and I wel-
come this opportunity of expressing my deep appreciation for
their assistance. My thanks go to: Mr. J. C. Battersby of the
British Museum for his kindness in answering sundry questions.
Dr. 0. G. S. Davis of Purdue University for allowing me to see
her manuscript synonymy of Python and Eryx. Dr. V. F.
FitzSimons of the Transvaal Museum for information regarding
certain South African amphibia, and early type localities not
to be found on recent maps. Dr. Carl Gans of Harvard Uni-
versity for rectifying my concept of the genus Dasypeltis and
permitting me to use his amended synonymy. Mons. Jean Guibe
of the Museum National d'Histoire Naturelle for examining the
Kenya specimen of Biifo latifrons. Henry Horn of Cambridge
High and Latin School for his conscientious and painstaking
work in checking citations, arranging the vast index, and related
matters. C. J. P. lonides, Esq., of Tanganyika Territory, whose

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 163

fine field work has added several new species to East Africa's
herpetofauna, and extended the known ranges of others. Miss
J. B. MacKenzie and her staff in the library of the Museum
of Comparative Zoology, for their ever willing help and skill in
tracking down references to obscure or inadequately cited pub-
lications. Dr. Ernst Mayr of this museum for advice on various
knotty problems of a taxonomic nature. Dr. Robert Mertens of
Senckenberg Museum for answering questions respecting early
German publications and Boettger types. Dr. H. W. Parker of
the British Museum for sundry information, and for pin-pointing
obscure localities in Ethiopia and Somalia. Mr. Benjamin
Shreve, my colleague for the past twenty-five years, for friendly
advice as to the gender and correct suffixes of sundry scientific
names. Dr. M. A. Smith of the British Museum for scanning
the pages relating to the synonymy of the HYDROPHIIDAE.
Dr. E. E. Williams of Harvard University for examining the
dentition of Cacostenium. and other favours. It is from our
recently published revision of the CRYPTODIRA that the
synonymy of the East African members of that suborder have
been taken. Miss N. E. Wright of this museum for her pains-
taking editorship of this manuscript.

Class REPTILIA

Subclass ANAPSIDA

Older TESTUDINATA'^

Suborder CRYPTODIRA"

Family DERMOCHELYIDAE

Genus DERMOCHELYS Blainville

1816. Dermocheh/s Blainville, Bull. Soc. Philom. Paris, p. "Ill" (mis-

6 The oldest name for the entire group is TESTUDINES Batsch, 1788 (Anleit.
Kennt. Thiere Mineral, 1, p. 437), rejected as being merely a nominative plural of
Testudo. f'HELONIA Macartney, 1802 (in Ross, Transl. Cuvier, List. Comp.
Anat. 1, pi. iii) is discarded as likely to result in confusion with the .crenus
Chelonia and with adjectives derived therefrom. Consequently, as the laws of
priority are not applicable to ordinal names. TESTUDINATA Oppel. 1811 (Ordn.
Kept., i). o) is preferred.

" Other suborders of TESTUDINATA are used by some herpetologists. On
account of its peculiar carapacial structure they would isolate Dermochelys as
suborder ATHECAE, all other turtles and tortoises being grouped under
THECOPHORA. Such a separation has been considered undesirable by some
anatomists and other recent workers.

164 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

print = 119). Type by monotypy: Testudo coriacea Linnaeus.
1820. Sphargis Merrem, Vers. Syst. Amphib., p. 19. Type by monotypy:

Testudo coriacea Linnaeus.
1822. Coriudo Fleming, Phil. Zool., 2, p. 271. Type by monotypy: Tes-

tudo coriacea Linnaeus.

1828. Seytina Wagler, Isis von Oken, 21, col. 861. Substitute name for

Sphargis Merrem.

1830. Dermatoclielys Wagler, Nat. Syst. Amphib., p. 133. Emendation
for Dcrmochelys.

1832. Chelyra Rafinesque, Atlantic Journ., 1. p. 64. Type by monotypy:
Testudo coriacea Linnaeus.
Dermochelys coriacea (Linnaeus) Luth or Leathery Turtle

17G(J. Testudo coriacea Linnaeus, Syst. Nat., ed. 12, 1, p. 350: Mediter-
ranean Sea.

1771. Testudo arcuata Catesby, Nat. Hist. Carolina, 2. p. 40: Coasts of
Carolina and Florida (by inference).

1788. Testudo lyra Lacepede, Hist. Nat. Quad. ovip. Serpens, 1. Synopsis
Methodiea (also p. Ill as Le Luth) : Barbary Coast, North
Africa.

1792. Testudo tuberculata Schoepff, Hist. Testud., p. 144: No locality.

1814. Chelonia lutaria Eafinesque, Specchio Sci. (Palermo), 2, No. 9, p.
66 : Mediterranean Sea.

1820. Sphargis mercuralis Merrem, Vers. Syst. Amphib., p. 19: Mediter-
ranean Sea and Atlantic Ocean.

1829. Dermochelis Atlantica Lcsueur {nomen nudum), in Cuvier, Regne

Animal, ed. 2, 2. p. 14, footnote : No locality.

1830. Dermatoclielys porcata Wagler, Nat. Syst. Amphib., pi. i, figs. 1-23

(on p. 133 coriacea is used) : No locality.
1884. Sphargis coriacea var. Sdhlegelii Garman, U. S. Nat. Mus. Bull.

No. 25, pp. 295, 303 : Tropical Pacific and Indian Oceans.
1899. Sphargis angusta Philippi, Anales Univ. Chile (Santiago), No.

104, p. 730, pi. i: Tocopilla, Chile.
Range. Indian and other tropical oceans, and, as an accidental
visitor, the temperate seas.

Family CHELONIIDAE
Genus CHELONIA Brongniart

1800. Chelonia Brongniart (part), Bull. Soe. Philom. Paris, 2. p. 89.
Type designated by Fitzinger:1843: Testudo mydas Linnaeus.

1837. Chelana Buraieister, Handbuch Naturg., 2. Abt. Zool., p. 731. Type

by monotypy: Testudo mydas Linnaeus.

1838. Mydas Cocteau (not Fabricius:1799), Rept., in Sagra, Hist. Fis.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 165

Pol. Xat. Cuba, 4. p. 22. Type by tautonomy: Testudo mydas

Linnaeus.
1843. Mydasea Gervais, Diet. Hist. Nat., 3. p. 457. Type by tautonomy:

Testudo mydas Linnaeus.
1845. Euchelonia Tschudi, Fauua Peruana, Herp., p. 22. Type by mono-

typy: Testudo mydas Linnaeus.
1848. Megemys Gistel, Naturg. Thierr., p. viii. Substitute name for

Chelonia Brongniart.
18.58. Euchelys Girard, U. S. Explor. Exped. 1838-1842, Herp-, p. 447.

TjT)e by monotypy: E. macropus Girard = Testudo mydas

Linnaeus.
1862. Chclone Strauch, Mem. Acad. Imp. Sci. St.-Petersbourg, (7) 5.

No. 7, p. 59. Type by original designation: Testudo viridis

Schneider = T. mydas Linnaeus.

Chelonia mydas (Liunaeus) Green Turtle
1758. Testudo mydas Linnaeus, Syst. Nat., ed. 10, 1, p. 197: Ascension
Island.

1782. Testudo (macropus) Walbaum, Chelongr. Schildkroten, p. 112: No

locality.

1783. Testudo Viridis Schneider, Natur. Schildkroten, p. 299: No

locality.

1787. Testudo japonica Thunberg, Vetensk. Acad. Handl., 8. p. 178, pi.

vii, fig. 1 : Japan.

1788. Testudo viarina (seu vulgaris) Lacepede, Hist. Nat. Quad. ovip.

Serpens, 1, Sj-nopsis Methodica (also p. 54 as La Tortue

Franche) : Torrid Zone.
1788. Testudo viridis-squamosa Lacepede, Hist. Nat. Quad. ovip. Serpens,

1, Synopsis Methodica (also p. 92 as La Tortue ecaille-verte) :

Amazon River, Brazil.
1802. Testudo cepedia-na Daudin, Hist. Nat. Rept., 2, p. 50, pi. xvii,

fig. 1 : No locality.
1812. Chelonia virgata Schweigger, Konigsberger Arch. Naturwiss.

Math., 1. pp. 291, 334, 411: Seas of the Torrid Zone.
1820. Caretta esculenta Merrem, Vers. Syst. Amphib., p. 18: Atlantic

Ocean.
1820. Caretta nasicornis Merrem, Vers. Syst. Amphib., p. 18: Ocean

near America.
1820. Caretta Thunbergii Merrem, Vers. Syst. Amphib., p. 19: Japan.
1829. Chelonia maculosa Cuvier, R^gne Animal, ed. 2, 2, p. 13: No local-
ity.
1829. Chelonia lachrymata Cuvier, RSgne Animal, ed. 2, 2, p. 13: No

locality.
1834. Chelonia bicarinata Lesson, in Belanger, Voy. Indies-Orient., Zool.,

p. 301 : Atlantic Ocean.

166 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1835. Chelonia Marmorata Dumeril & Bibron, Erpet. Gen., 2, p. 546, pi.

xxiii, fig. 1 : Ascension Island.
1858. Chelonia formosa Girard, U. S. Explor. Exped. 1838-1842, Herp.,

p. 456, pi. xxxi, figs. 1-4: Fiji Islands.
1858. Chcloniu tenuis Girard, U. S. Explor. Exped. 1838-1842, Herp.,

p. 459, pi. xx.xi, fig. 8: Honden Island, Paumotu Group, Tahiti

and Eimeo ; Rosa Island.
1868. Chelonia Agassisii Bocourt, Ann. Sci. Nat. Zool. (Paris), (5) 10,

p. 122: Nagulate River mouth, Guatemala.
1880. Chelonia depressa Garman (at least in part), Bull. Mus. Comp.

Zool., 6, p. 124: East Indies (juvenile).
1887. Chelonia lata Philippi, Zool. Garten, 28. p 84: near Valparaiso,

Chile.
Range. Indian and other tropical oceans and, as an accidental
visitor, the temperate seas.

Genus ERETMOCHELYS Fitzinger

1828. Carctta Ritgen (not Rafinesque:1814), Nova Acta Acad. Leop.-
Carol., 14. p. 270. Type by monotopy: Testudo imhricata
Linnaeus.

1843. Eretmochelys Fitzinger, Syst. Rept., p. 30. Type by original desig-
nation: Testudo imbricata Linnaeus.

1868. Herpysmostes Gistel in Blicke, Leben Natur Menschen (Leipzig),
p. 145. Type (fide Mertens:1936) : Testudo imbricata Linnaeus.

1873. Onychochelys Gray, Proc. Zool. Soc. London, p. 397, figs. 1-2. Type
by monotypy: 0. Icraussi Gray = Testudo imbricata Linnaeus.

Eretmochelys imbricata (Linnaeus) Hawksbill Turtle

1766. Testudo imbricata Linnaeus, Syst. Nat., ed. 12, 1. p. 3r)0: American

and Asiatic seas.
1834. Chelonia psevdo-mydas Lesson, in Belanger, Voy. Indies-Orient.,

Zool., p. 299 : Atlantic Ocean.

1834. Chelonia pseudo-caretta Lesson, in Belanger, Voy. Indies-Orient..

Zool., p. 302 : Atlantic Ocean.

1835. Caretta Bissa Riippell, Neue Wirbelth. Fauna Abyssinica, Amphib.,

p. 4, pi. ii: Red Sea.

1857. Eretmochelys squamata Agassiz, Contr. Nat. Hist. U. S., 1. p. 382:

Indian and Pacific Oceans.

1858. Caretta squamosa Girard, U. S. Explor. Exped. 1838-1842, Herp.,

p. 442, pi. XXX, figs. 1-7: Sulu Seas and Indian Ocean.
1858. Caretta rostrata Girard, U. S. Explor. Exped. 1838-1842, Herp.,

p. 446, pi. XXX, figs. 8-13: Fiji Islands.
1873. Onychochelys Icraussi Gray, Proc. Zool. Soc. London, p. 398, figs.

1-2: Atlantic Ocean off French Guiana.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 167

Range. Indian and other tropical oceans and, as an accidental
visitor, the temperate seas.

Genus CARETTA Rafinesque

181-1. Caretta Rafinesque, Speechio Sci. (Palermo), 2, No. 9, p. 66. Type

by monotypy: C. nasuta Rafinesque = Testudo caretta Linnaeus.
1835. Thalassochelys Fitzinger, Ann. Wiener Mus., 1, pp, 110, 121, 128.

Type by original designation: T. caouana Fitzinger = Testudo

caretta Linnaeus.
1838. Caouana Cocteau, Rept., in Sagra, Hist. Fis. Pol. Nat. Cuba, 4. p.

31. Type by monotypy: Thalassochelys cephalo Schneider :=

Testudo caretta Linnaeus.

1843. UaUchelys Fitzinger, Syst. Rept., p. 30. Type by original designa-

tion: H. atra Fitzinger = Testudo caretta Linnaeus.

1873. Cephalochelys Gray, Proc. Zool. Soc. London, p. 408. Type by
monotypy: C. oceanica Gray = Testudo caretta Linnaeus.

1873. Eremonia Gray, Proc. Zool. Soc. London, p. 408. Type by mono-
typy: E. elongata Gray =^ Testudo caretta Linnaeus.

Caretta caretta (Linnaeus) Red-brown Loggerhead

1758. Testudo Caretta Linnaeus, Syst. Nat., ed. 10, 1. p. 197: Islands

off America.
1783. Testudo Cephalo Schneider, Natur. Schildkroten, p. 303: No

locality.
1788. Testudo camia7ia Lacepede, Hist. Nat. Quad. ovip. Serpens, I,

Synopsis Methodica (also p. 95 as La Caouane) : Jamaica, British

West Indies (restricted).
1788. Testudo nasicornis Lacepede, Hist. Nat; Quad. ovip. Serpens, 1,

Synopsis Methodica (also p. 103 as La Tortue nasicorne) :

America.
1814. Caretta nasuta Rafinesque, Speechio Sci. (Palermo), 2. No. 9,

p. 66: Sicily (fide Lindholm:1929).
1820. Caretta atra Merrem, Vers. Syst. Amphib., p. 17: Ascension

Island.
1833. Chelonia pelasgorum Valenciennes, pi. vi, in Bory, Exped. Sci.

Moree, Zool.: Modon, etc., Mediterranean Sea.

1844. Caouana elongata Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 53:

No locality.
1858. Thalassochelys corticata Girard, U. S. Explor. Exped. 1838-1842,

Herp., p. 431, pi. xxix: Madeira Islands.
1873. Cephalochelys oceanica Gray, Proc. Zool. Soc. London, p. 408:

Mexico.
1887. Thalassochelys tarapacona (sic) Philippi (identified by subsequent

1 08 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

description of Philippi:1899), Zool. Garten, 28, p. 83: Iquique,
Chile.
,1930. Caretia gigas Deraniyagala, Ceylon Journ. Sci., (B), 18, p. 66,
figs. 4-6, pi. v: Gulf of Mannar, Ceylon.
Range. Indian and other tropical oceans and, as an accidental
visitor, the temperate seas.

Genus LEPIDOCHELYS Fitzinger

1843. Lepidochelys Fitzinger, Syst. Eept., p. 30. Type by original desig-

nation: Chelonia olivacea Eschscholtz.
1880. Colpochelys Garman, Bull. Mus. Coiup. Zool., 8, p. 124. Type by
monotypy : C. Tcempi Garman.

Lepidochelys oUvacea olivacea (Eschscholtz)
Olive Loggerhead
1820. Whelonia multisnitata Kuhl, Beitr, Zool. Anat., 1. p. 78: No

locality.
1829. Chelonia olivacea Eschscholtz, Zool. Atlas, p. 3, pi. iii: Manila Bay,

Philippine Islands.
1835. Chelonia Bussumierii Dumeril & Bibron, Erpet. Gen., 2, p. 557, pi.
xxiv, figs. 1-la: China Sea and Malabar Coast.

1844. Caoiuvna Riippcllii Gray {nomen nudum). Cat. Tort. Croc. Amphis.

Brit. Mus., p. 53: India?

1857. Chelonia polyaspis Bleeker, Nat. Tijdschr. Ned. Indies, 14, p. 239:
Batavia, Java.

1889. Chelonia duhia Bleeker {nomen nudum), in Boulenger, Cat. Chelon.
Ehj'ncho. Croc. Brit. Mus., p. 186: Borneo.

1899. TJialassochelys controversa Philippi, Anales Univ. Chile (Santi-
ago), No. 104, p. 732: No locality.

1908. Caretta remivaga Hay, Proc. U. S. Nat. Mus., 34. p. 194, pi. x,
figs. 1-3 ; pi. xi, fig. 5 : Ventosa Bay, Gulf of Tehuantepec,
Mexico.

Range. Indian and other tropical oceans and, as an accidental
visitor, the temperate seas.

Family TESTUDINIDAE
Genus TESTUDO Linnaeus^

1758. Testudo Linnaeus, Syst. Nat., ed. 10, 1, p. 197. Type by subsequent

designation of Fitzinger: Testudo graeea Linnaeus.

s In place of the single genus Testudo some authorities would recognize several
full genera {Testudo; Geochelonej Psammobates) . Despite possibly ancient sep-
aration I personally prefer to regard these groups as of only subgeneric rank.
However, this is merely my personal opinion ; for those who think otherwise
and consider Geochelone as a full genus, the only synonyms of it would be
Centrochelysj Stigmochelys and Megaohersine.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 169

1820. Chersine Merrem, Vers. Syst. Amphib., pp. 12, 29. Type by subse

quent designation of Lindholm: Testudo graeca Linnaeus.
1830. Chersus Wagler, Nat. Syst. Amphib., p. 138. TjTje by monotypy:

Testudo marginata Schoepff.
1835. Psammobates Fitzinger, Ann. Weiner Mus., 1, pp. 108, 113, 122.

Type by original designation: Testudo geometrica Linnaeus.
1835. Geochelone Fitzinger, Ann. Weiner Mus., 1, pp. Ill, 112, 122.

Type by original designation: Testudo stellata Schweigger = T.

clegans Schoepff.

1869. Peltastes Gray (not Bossi:1807), Proc. Zool. Soc. London, pp.

167, 171. Type by subsequent designation of Lindholm: Testudo
graeca Linnaeus.

1870. Chersin.ella Gray, Suppl. Cat. Shield Rept. Brit. Mus., part 1, p. 8.

Type by subsequent designation of Lindholm: Testudo graeca

Linnaeus.
1870. Testudinella Gray (not Bory:1826), Suppl. Cat, Shield Rept. Brit.

Mus., part 1, p. 12. Type by monotypy: Testudo horsfieldi Gray.
1872. Peltonia Gray, App. Cat. Shield Rept. Brit. Mus., p. 4. New name

for Peltastes Gray (preoccupied).

1872. Centrochelys Gray, App. Cat. Shield Rept. Brit. Mus., p. 5. Type

by monotypy: Testudo sulcata Miller.

1873. Stigmochelys Gray, Hand-List Shield Rept. Brit. Mus., p. 5. Type

by monotypy : Testudo pardalis Bell.
1916. Medaestia Wussow, Wochenschr. Aquar.-Terrar. Kunde, 13, p. 170.

Type by subsequent designation of Mertens: Testudo graeca

Linnaeus.
1933. Megachersine Hewitt, Ann. Natal Mus., 7, p. 257. Type by original

designation : Testudo pardalis Bell.
1933. Chersinella Hewitt (not Gray: 1870), Ann. Natal Mus., 7, p. 259.

Type by original designation: Testudo geometrica Linnaeus.

Testudo pardalis babcocki Loveridge Eastern Leopard Tortoise
1935. Testudo pardalis hahcocki Loveridge, Bull. Mus. Comp. Zool., 79.

p. 4: Mount Debasien, Karamojo, Uganda.
Range. Sudan and Ethiopia, south through Uganda; Kenya
Colony and Tanganyika Territory to Natal, west through Cape
Province to Southwest Africa where it meets with the typical
form, but is dominant north of 25° S. and in southern Angola.

Genus MALACOCHERSUS Lindholm

1929. Malacochersus Lindholm, Zool. Anz., 81, p. 285. Type by original
designation : Testudo tornieri Siebenrock.
Molaco chersus tornieri (Siebenrock) Pancake Tortoise

1 70 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1903. Testudo tornicri Siebeiirock, Auz. Akad. Wiss. Wien, 40, p. 185 :

Busisi, Smith Sound, Lake Victoria, Tanganyika Territory.
1920. Tcstndo Lovcridgii Boulenger, C. R. Acad. Sci. (Paris), 170, p. 263:

Dodonia, Ugogo, Tanganyika Territory.
1923. '^Testudo procterae Loveridge, Proc. Zool. Soc. London, p. 298, pis.

i-ii: Ikikuyu, south of Gulwe, Tanganyika Territory."
Range. Rocky hills of Kenya Colony (Mathews Range south
to Njoro. east to Midas Creek), and Tanganyika Territorj^ (fi'oni
Busisi, Smith Sound, southeast through Ugogo to Lindi).

Genus KINIXYS Bell

1827. Kinixys Bell, Trans. Linn. Soc. Loudon, 15, p. 398. Type by

original designation : K. castanea Bell = Testudo erosa Schweig-

ger.
1835. CinotJiorax Fitzinger, Ann. Wiener Mus., 1, pp. 108, 111, 121. Type

by subsequent designation of Fitzinger : 1843 : Kinixys helliana

Gray.
1843. Cinixys Fitzinger, Syst. Kept., p. 29. Emendation for Kinixys

Bell.
1873. Klnothorax Gray, Hand-List Shield Eept. Brit. Mus., p. 16.

Emendation for CinotJiorax Fitzinger.

Kinixys belliana belliana Gray Bell's Eastern Hinged-Tortoise
1831. Kinixys Belliana Gray, Synopsis Reptilium, p. 69: No locality.

("W. Africa ?", later added by Boulenger, appears doubtful.)
1845. Kinixys sehoensis Riippell, Mus. Senckenberg., 3, p. 226, pi. xvi,

figs. 1-3 : Shoa, southern Ethiopia.
1863. Kinixys Spcl'ti Gray, Ann. Mag. Nat. Hist., (3) 12, p. 381:

"Central Africa" (probably northwest Tanganyika Territory).
1902. Homopus darlingi Boulenger, Proc. Zool. Soc, London, 2, p, 15,

pi. iv: Salisbury District, Mashonaland, Southern Rhodesia,
1927. Cinixys lohatsiana Power, Trans. Roy. Soc. S. Africa, 14. p, 410,

pis. xix-xx: Lobatsi, Bechuanaland Protectorate.
1931. Kinixys helliana sombensis Hewitt, Ann. Natal Mus., 6, p. 469, fig.

Ic, pi. xxxviii, fig, 4: Zomba, Nyasaland,
1931. Kinixys belliana zuluensis Hewitt, Ann, Natal Mus., 6, p. 471, figs.

la-lb, pi. xxxviii, figs. 1-3, 5-10: Richard's Bay, Zululand,
1931. Kinixys austraUs Hewitt, Ann. Natal Mus., 6, p. 477, pi, xxxvi,

figs. 4-6 : White River, Nelspruit, Lydenburg District, Transvaal.

9 As Procter's Soft-shelled Land-Tortoise is known only from a single specimen,
it is tentatively relegated to tlie synonymy on geographical grounds until someone
can secure a series from tlie type locality, which should not be confused with
the Ikikuyu just outside Dodoma. Not only is M. procterae strikingly different in
coloration from every one of the more than a hundred tornicri I have seen, but
also its head scalation, especially its large and elongate prefrontals, is unique.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 171

1931. Kinixya jordani Hewitt, Ann. Natal Mus., 6. p. 481, pi. xxxvii, figs.
7-9 {not 1-3 as stated): Isoka, Northern Rhodesia.

1931. Kvnixys youngi Hewitt, Ann. Natal Mus., 6. p. 486, fig. Id, pi.

xxxvii. figs. 4-5: Lake Nyasa shore below Livingstonia, Nyasa-
land.

1932. Kinixys australis mababiensis FitzSimons, Ann. Transvaal Mus.,

15. p. 37: Tsotsoroga Pan, Mababe Flats, Bechuanaland Pro
tectorate.
1935. Kinixys natalensis Hewitt, Eee. Albany Mus., 4, p. 353, pi. xxxv,

figs. 3-4: Jameson's Drift, Tugela River Valley, Natal.
Range. Eritrea southwest to Natal, northwest through Bechu-
analand and Angola to French Cameroon where it meets with
the western race (which has only four claws on forelimb).
Madagascar.

Kinixys erosa (Schweigger) Western-Forest Hinged-Tortoise
1812. Tcstudo erosa Schweigger, Konigsberger Arch. Naturwiss. Math..
I. p. 321: "America septentrionali" (error).

1826. Testiido Schopfii Fitzinger (nomen nudum), Neue Class. Rept., p.

44 : No locality.

1827. Kinixys castanca Bell, Trans. Linn. Soc. London, 15. p. 398, pi.

xvii, fig. 1 : Africa.
Range. Uganda, west through the Belgian Congo to Gambia.

Family TRIONYCHIDAE
Genus TRIONYX Geoffrey '-^

1809. Trionyx Geoffroy Saint-Hilaire, Ann. Mus. Hist. Nat. (Paris),
14. p. 1. Type by original designation: T. aegyptiacus Geoffroy
= Testudo iriunguis Forskal.

1830. Aspidonectcs Wagler, Nat. Syst. Amphib., p. 134. Type by desig-
nation of Stejneger: Trionyx aegyptiacus Geoffroy = Testudo
triunguis Forsk°al.

1835. Gymnopus Dumeril & Bibron, Erpet. Gen., 2, p. 472. New name
for Aspidonectes Wagler.

1844. Tyrse Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 47. Type by
tautonomy: Trionyx nilotica Gray = Testudo triunguis Forskal.

1869. Fordia Gray, Proc. Zool. Soc. London, p. 219. Type by monotypy:
F. africana Gray = Testudo triimguis Forskal.

10 Only synonyms based on African species are given here as it seems scarcely
v.'orth wliile to list the 24 additional ones involving American and Asiatic species.
These can be found in the recently published revision of African Cryptodira (see
Bibliography).

172 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Trionyx triunguis (Forskal) Nile Soft-shelled Turtle

1775. Testudo triunguis Forskal, Descr. Anim. Avium, Amphib., p. ix:

Nile, Egypt.
1809. Trionyx Aegyptiacus Geoffroy Saint-Hilaire, Ann. Mus. Hist. Nat.

(Paris), 14, p. 12, pis. i-ii: Nile, Egypt.
1831. Trionyx Niloticus "Shaw" Gray, Synopsis Reptilium, p. 46:

Nile, Egyi:)t.
1837. Trionyx lahiatus Bell, Monog. Testudinata, text to pis. — (=

xviii-xx) : Sierra Leone.
1844. Trionyx Mortoni Hallow ell, Proc. Acad. Nat. Sci. Philadelphia, p.

120: Africa.
1859. Aspidonectes aspilus Cope, Proc. Acad. Nat. Sci. Philadelphia, p.

295 : Ovenga and Rembo Rivers, Fernan Vaz, French Congo.
1869. Fordia africana Gray, Proc. Zool. Soc. London, p. 219: Nile River

at Khartoum, Sudan.
1948. Trionyx triunguis rudolfianus Deraniyagala, Spolia Zeylanica

(Colombo), 25, 2, p. 30, fig. 5, pi. xii, fig. c: Ferguson's Gulf,

Lake Rudolf, Kenya Colony.
Range. Egypt and Eritrea south to Lake Rudolf, Kenya Col-
ony; southwest to Lake Albert, Uganda; and Angola; northwest
to Senegal. Accidental visitor along the Palestine coast of Asia
Minor.

Genus CYCLODERMA Peters

1854. Cycloderma Peters, Monatsb. Akad. Wiss. Berlin, p. 216. Type

by monotypy : C. frenatum Peters.
1856. Cryptopodiis A. Dum^ril, Revue Mag. Zool. (2), 8, p. 374. Lapsus

for Cryptopus Dumeril & Bibron = Cyclanorbis (part).

1859. Heptathyra Cope, Proc. Nat. Sci. Philadelphia, p. 294. Type by

monotypy: Cryptopus (sic) aubryi A. Dumeril,

1860. Aspidochelys Gray, Proc. Zool. Soc. London, p. 6. Type by mono

typy: A. livingstonii Gray := Cycloderma frenatum Peters.

Cycloderma frenatum Peters Zambezi Soft-shelled Turtle

1854. Cycloderma frenatum Peters, Monatsb, Akad. Wiss. Berlin, p. 216:

Zambezi River, Mozambique.
I860. Aspidochelys livingstonii Gray, Proc. Zool. Soc. London, p. 6, pi.

xxii, figs. 1-2: Tributaries of Zambezi, Mozambique.
Range. Southern Tanganyika Territory, west and south
through Nyasaland (and possibly Northern Rhodesia) to Mozam-
bique.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 17:)

Suborder PLEURODIRA

Family PELOMEDUSIDAE

Genus PELOMEDUSA Wagler

1830. Pelomedusa Wagler, Nat. Syst. Amphib., p. 13G. Type by iiiono-
typy: Testudo galeata Schoepff = Testudo suhrufa Lacepede.

1835. Pentonyx Dum^ril & Bibron, Erpet. Gen., 2. p. 389. Type by
subsequent designation: P. capensis Dumeril & Bibron =r Tes-
tudo stibrufa Lacepede.

Pelomedusa subruia olivacea (Seliweigger)"
Northeru Marsh-Terrapin
1814. Emys olivacea Schweigger, Prodromi Mon. Chelon., p. 38: "In

Fabulosis Nigritae," as Adanson coll. = Senegal.
1835. Pentonyx G.ehafie RiippcU, Neue Wirbelth. Fauna Abyssinica,

Amphib., p. 2, pi. 1 : Massaua, Eritrea.
1884. Pelomedusa Gasconi Rochebrune, Faune Senegambie, Eept., p. 25,

pi. i, figs. 1-2: Dagana, Senegal (restricted).
1910. Welomedusa galeata var. disjuncta Vaillant & Grandidier, Hist.
Phys. Nat. Pol. Madagascar, 17. Eept., p. 56, pi. xx, fig. 3: Du
Bourg de Bozas coll., but no locality.
Range. Uncertain, but apparently a northern belt extending
from Senegal to Eritrea and intergrading with the typical form
in northern Uganda and northern Kenya Colony with occasional
indi\aduals cropping uj) in the south and Madagascar.

Pelomedusa subruia subruia (Lacepede)
Southern Marsh-Terrapin
1788. Testudo suhrufa Lacepede, Hist. Nat. Quad. ovip. Serpens, 1,

Synopsis Methodica (also p. 173 as La Eoussatre): "de

L'Inde, " as collected by Sonnerat restricted to Cape of Good

Hope, South Africa.
1792. Testudo galeata Schoepff, Hist. Testud., p. 12, pi. iii, fig. 1:

"India orientale, Carolina," restricted to Cape Flats, Cape

Province, South Africa.
1798. Testudo Badia Donndorf, Zool. Beytrage Linn. Natur., 3, p. 34

(new name for Lacepede 's La Eoussatre): No locality.
1835. Pentonyx Capensis Dumeril & Bibron, Erpet. Gen., 2, p. 390, pi.

11 Some authorities would relegate this race and its forms to the synonymy
of subruja. It is separated here in order to stimulate inquiry as to whether or
not 75 per cent of the northern population have their pectoral shields separated
on the median line of the plastron — the sole character allegedly separating the
two races. P. g. disjuncta, if indeed it actually came from Madagascar, may be a
throwbacit for these do occur, though very rarely, within the range of the typical
s. subrufa.

174 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

xix, figs. 2-2b : restricted by Mertens to Cape of Good Hope,
South Africa.
1849. •Pentonix americana Cornalia, Vert. Synop. Mus. Mediolau, p. 13:
"in flumine propre Novaeboracum, " i.e. New York (error).

1855. Pelomedusa Mozamhica "Peters M.S.S. 1848" Gray (Nomen

nudutn) , Cat. Shield Rept. Brit. Mus., p. 53: Mozambique.

1856. Felomedusa mossambicensis "Peters" Lichtenstein & v. Martens,

{nomen nudum) Nomencl. Eept., p. 2 : Mozambique.
1863. Pelomedusa nigra Gray, Ann. Mag. Nat. Hist., (3) 12, p. 99:

Natal, South Africa.
1910. IPelomedusa galeata var. disjuncta VaiUant & Grandidier, Hist.

Phys. Nat. Pol. Madagascar, 17, p. 56: Madagascar.^-
1935. Pelomedusa galeata orangensis Hewitt, Rec. Albany Mus., 4, p.

332, pi. xxxi, fig. 3; pi. xxxii, figs. 3-4: Kimberley, Cape Prov-
ince, South Africa.
1935. Pelomedusa galeata devilliersi Hewitt, Rec. Albany Mus., 4, p. 337,

pi. xxxi, figs. 2 and 4: Besondermeid, Steinkopf, Cape Province,

South Africa.
1935. Pelomedusa galeata damarensis Hewitt, Rec. Albany Mus., 4, p.

338, pi. xxxiii, figs. 1-4: Quickborn Farm, near Okahandja,

Southwest Africa.
1937. Pelomedusa subrufa wettstcini Mertens, Zool. Anz., 117, p. 141,

figs. 1 and 4: Majunga, western Madagascar.
Range. Dry savanna of southern Sudan east to British Somali-
land, south through Uganda;^" Kenya Colony and Tanganyika
Territory to Natal, west through Cape Province to Southwest
Africa, north and northeast to Belgian Congo.

Genus PELUSIOS Wagler

1830. Pelusios Wagler, Nat. Syst. Amphib., p. 137. Type by monotypy :
Einys eastanea Sehweigger = Testudo subnigra Lacepdde.

18G3. Tanoa Gray, Proe. Zool. Soc. London, p. 193. Type by monotypy:
Sternothaerus siiiuatus A. Smith.

1863. Notoa Gray, Proc. Zool. Soc. London, p. 195. Type by monotypy:
Sternothaerus suhniger Gray = Testudo subnigra Lacepede.

Pelusios subniger (Lacepede)^" Black Terrapin

1 7SS. Testudo subnigra Lacepede, Hist. Nat. Quad. ovip. Serpens, I,
Synopsis Methodica (also p. 175, pi. xiii as La Noiratre): No
locality.

12 But see note under Pelomedusa s. olivacea (Sehweigger).

13 From this s.vnonymy I have removed f^tcrnothcrus niger Dum6ril & Bibron,
together with its synonym .Sf. oxiirhuniD Boulenper, as they represent a valid
West Afriean siioeies.

LOVERJDGE : E. AFRICAN REPTILES AND AMPHIBIANS 17."i

1798. Testudo nigricans Donndorff, Zool. Beytrage Linn. Natur. 3, p. 34
(new name for La Noiratre Lacepede) : No locality.

1814. Emi/s castanea Scliweigger, Prodromi Men. Chelou., p. 45 (new
name for siihniffra Daudin in Paris Museum) : No locality.

1825. Sternotltaerus Leachinnus Bell, Zool. Journ., 2, p. 306: No locality.

1844. Sternotherns Dcrbianus Gray, Cat. Tort. Croc. Amphis. Brit. Mus.,
p. 37: Gambia (restricted).

1851. Sternotlierus nigrescens Bianconi (lapsus for nigricans used in the
text). Spec. Zool. Mosamb., Kept., pi. vii.

1886. Pelomedusa Jouberti Eochebrune, Vertebrata nov. Africae Occi-
dent., Part 3, p. 10: Noki, Congo Eiver, Angola; Landana,
Cabinda ; and Mellacoree Eiver, French Guinea.

1906. Sternothaeriis nigricans seycliellcnsis Siebenrock, in Voeltzkow,
Eeise in Ostafrika, 2. p. 38: Gloriosa Island.

1927. Peliisios nigricans rhodesianns Hewitt, Eec. Albany Mus., 4. p.
375, figs, la, Ic ; pi. xxvi. figs. 2-3: Mpika District, Northern
Ehodesia.

1931. Peliisios nigricans castanoides Hewitt, Ann. Natal Mus., 6. p. 463,

pi. xxxvi, figs. 1-2: Eichard 's Bay, Zululand.

1932. Pelusios bechuanicus FitzSimous, Ann. Transvaal, Mus., 15, p. 37:

Thamalakaue Eiver at Maun, Bechuanaland Protectorate.
Range. Uganda ; Kenya Colony ; Tanganyika Territory and
islands of the Indian Ocean (Pemba; Zanzibar; Seychelles;
Madagascar; Mauritius). In east, south to Zululand, Natal,
northwest to Angola ; Senegal and Cape A'erde Islands, east to
T'ganda.

Pelusios sinuatus (Smith) Serrated Terrapin

ls38. Sternotherns sinuatus A. Smith, lUus. Zool. S. Africa, Eept., pi. i:

In rivers to the north of 25° S., South Africa.
1848. Sternotherns dentatus Peters, Arch. Anat. Phys., p. 494: No

locality.
1895. Sternotltaerus bottegi Boulenger, Ann. Mus. Civ. Stor. Nat. Genova,

(2) 15. p. 9, pis. i-ii: Bardera, Somalia.
1927. Pelusios sinuatus suluensis Hewitt, Eec. Albany Mus., 4, p. 360.

fig. Id, pi. XX, figs. 1-3: near Umsinene Eiver, Zululand.

1933. Pelusios sinuatus leptus Hewitt, Occ. Papers Ehodesian Mus., p. 45,

jjI. ix, figs. 1-2: Isoka, Northern Ehodesia.
Range. Somalia, south through Kenya Colony and Tanganyika
Territory to Natal, northwest to the eastern Belgian Congo."

1+ Unknown from Uganda and Lake Victoria, for early records from tlie Sessc
Islands and Hulcoba were based im misidentilied suhttigcr.

176 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Subclass ARCHOSAURIA

Order CROCODYLIA^'

Family CROCODYLIDAE

Genus CROCODYLUS Laurenti'''

1763. Crocodylus Gronovius, Zoophyl. Gronov., fasc. 1, p. 10. Rendered

unavailable by the Internat. Comm. on Zool. Nomen.^'^
1768. Crocodylus Laurenti, Syn. Rept., p. 53. Type by subsequent

designation of Stejneger & Barbour : 1917: C. niloticus Laurenti

= Lacerta crocodilus Linnaeus (part).
1789. Crocodilus Bonnaterre (part), Tabl. Encycl. Method. Regnes Nat.,

Erpet., p. 32. Emendation for Crocodylus Laurenti.
1820. Oiampse Merrem, Vers. Syst. Amphib., p. 36. Type by subsequent

designation: Crocodilus biscutatus Cuvier = C. acutus Cuvier.
1844. Mecistops Gray (part), Cat. Tort. Croc. Amphis. Brit. Mus. Type

by subsequent designation Crocodilus cataphractus Cuvier.
1844. Oopholis Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 58. Type

by subsequent designation of Deraniyagala: Crocodilus porosus

Schneider
1844. Palinia Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 60. Type by

monotypy: Crocodilus rhomhifer Cuvier.
1844. Motinia'^^ Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 60. Type

by monotypy: Crocodilus americanus Schneider = C. acutus

Cuvier.
1862. Bombifrons Gray, Ann. Mag. Nat. Hist., (3) 10. p. 269. Type by

subsequent designation: Crocodilus bombifrons Gray = C.

palustris Lesson.
1862. remsacus Gray, Ann. Mag. Nat. Hist., (3) 10, p. 272. Type by

monotypy : Crocodilus intermedius Graves.
1874. Philas Gray, Proc. Zool. Soe. London, p. 188. Type by monotypy:

Crocodilus johnsoni Krefft := C. johnstoni Krefft.

Crocodylus cataphractus Cuvier Long-nosed Crocodile

1801. ^Crocodilus )iiger Latreille, in Buffon, Hist. Nat. Rept, (ed. D6ter-
ville), 1, p. 210: Senegal River, Senegal.

15 Generally accepted by herpetologists in preference to LORICATA which lias
been applied to pangolins and other groups.

16 The skull from "Afrika ?" named Champse brevirostris (or Crocodilus
krevirostrig in the caption to figs.) Werner, 1933, Zool. Anz., 102, p. lOO, figs.
1-2, has been identified as the Asiatic Crocodylus palustris Lesson by VVermuth,
1953, Mitt. Zool. Mus. Berlin, 29, p. 472.

17 See Opinions and Declarations, 1926, in Smithsonian Misc. Coll., 73, pp. 27-33.

ISA misprint, corrected to Molinia by Gray, 18(52, Ann. Mag. Nat. Uist., (3) 10,
p. 272.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 177

1824. Crocodiliis cataphractm Cuvier, Rech. Ossem. foss. Quad., ed. 2,
5. p. 58, pi. V, figs. 1-2: No locality.

1835. Crocodilus leptorhynchus Bennett, Proc. Zool. Soc. London, p. 129:
Fernando Po.

1844. Mecistops Bennettii Gray (new name for C. Icptorhynchns Ben-
nett), Cat. Tort. Croc. Aniphis. Brit. Mus., p. 57.

Range. Near Ujiji, Lake Tanganyika, Tanganyika Territory,
west to Angola, northwest to Senegal.

Crocodylus niloticus Laurenti Nile Crocodile

1768. Crocodylus niloticus Laurenti (part), Syn. Kept., p. 53: Egypt
(restricted).

1768. Crocodylus africanus Laurenti (part; with part lizard, so unidenti-
fiable), Syn. Kept., p. 54: No locality.

1807. Crocodilus vulgaris Cuvier, Ann. Mus. Hist. Nat. (Paris), 10,
p. 40, pis. iiii : Egypt.

1807. Crocodilus suchus Geoffroy Saint-Hilairc, Ann. Mus. Hist. Nat.
(Paris), 10, p. 84, pi. iii: Nile and Niger Rivers.

1824. Crocodilus Chamses Bory de Saint-Vincent, Diet. Class. Hist. Nat.,
5. p. 105: Nile, Egypt.

1826. Crocodilus muULscutatus Riippell, in Cretzschmar, Iris (Frankfurt

am Main), 25. p. 99: Soucot, Nile, "Nubia" i.e. Kordofan,
Sudan.

1827. Crocodilus marginatus Geoffroy Saiut-Hilaire, Crocodiles, in Descr.

Egypte, p. 260: Nile River near Thebes, Egj-pt.
1827. Crocodilus lacunosus Geoffroy Saint-Hilaire, Crocodiles, in Descr.

Egypte, p. 261: (mummy) EgjT)t.
1827. Crocodilus complatiatus Geoffroy Saint-Hilaire, Crocodiles, in

Descr. Egypte, p. 263: (mummy) Egypt.
1831. Alligator cowieii A. Smith, S. African Quart. Journ., (5) p. 15:

"Rivers beyond Currichane, " i.e. Rustenburg, western Trans-
vaal.
1831. Crocodilus Octophractus Gray, in Griffith, Animal Kingdom, 9.

Synopsis, p. 22: Soucot, Nile River, Kordofan, Sudan.
1857. Crocodilus Binuensis Baikie, Proc. Zool. Soc. London, p. 48:

Benue and Niger Rivers, Nigeria.
1872. Crocodilus inadagascariensis Grandidier, Ann. Sci. Nat. Zool.

(Paris), 15. art. 20, p. 6: Madagascar.
1872. Crocodilus robustus Grandidier & Vaillant, C. R. Acad. Sci. (Paris),

75. p. 150: (fossil) Amboulintsatre, Madagascar.
1886. Crocodilus hexaphractos Ruppell {nomen nudum), in Schmidt, Ber.

Senckenberg. Naturf. Ges., for 1885, p. 131.
1948. Crocodylus niloticus loortliingtoni Deraniyagala, Spolia Zeylanica

(Colombo), 25, 2, p. 30: Lake Baringo, Kenya Colony.

178 BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY

1948. Crocodylus niloticus pauciscutatus Deraniyagala, Spolia Zeylanica
(Colombo), 25. 2, p. 31: Lake Rudolf, Kenya Colony.

Range. Uganda ; Kenya Colony ; Tanganyika Territorj^ : also
most lakes and rivers of Africa from about 20° N., south to
Tugela River, Natal. Zanzibar^'' and other islands (Seychelles;
Comoros; Madagascar) of the Indian Ocean; also present in the
Zerka River, Jordan.

Genus OSTEOLAEMUS Cope

1861. Osteolaemus Cope, Pioc. Acad. Nat. Sci. Philadelphia for I860,

p. 549. Type by monotypy: Osteolaemus tetraspis Cope.

1862. Halcrosia Gray, Ann. Mag. Nat. Hist., (3) 10. p. 273. Type by

monotypy: Crocodilus frontatus Alurray.
1919. Ostcohlepharon K. P. Schmidt, Bull. Amer. Mus. Nat. Hist., 39,
p. 420. Type by original designation : 0. osborni Schmidt.

Osteolaemus tetraspis osborni (Schmidt)
Eastern Broad-nosed Crocodile
1919. Osteohlepharon osborni K. P. Schmidt, Bull. Amer. Mus. Nat.
Hist., 39, p. 421, figs, 2-5, pi. xiii, fig. 1 : Niapu, Belgian Congo.
Range. Western Uganda and eastern Belgian Congo.'"

Subclass SYNAPTOSAURIA

Order SQUAMATA

Suborder SAURIA=^^

Family EUBLEPHARIDAE"-

Genus HOLODACTYLUS Boettger

1893. Holodactylus Boettger, Zool. Anz., 16. p. 113. Type by monotypy:
E. africanus Boettger.

Holodactylus africanus Boettger

1893. Holodactylus africanus Boettger, Zool. Anz., 16, p. 114: "Abdal-
lah," i.e. Abdulla, north of Webi Shebeli, Ethiopia.

ly As an accidental visitor : now mounted in the Peace Memorial Museum,
Zanzibar.

20 Fortunately the onl.v Uganda specimen was photographed prior to its escape.
The typical form (O. t. tetraspis Cope) ranges from the western Belgian Congo
and Angola, northwest to Senegal.

21 LACERTILIA of Owen (1842), Boulenger (1S85) and others.

^2 Geckos with true eyelids. This family, long merged with the GEKKONIDAE,
was revived by Underwood (1954, Proc. Zool. Soc. London, 124. p. 476).

LOVERIDQE: E. AFRICAN REPTILES AND AMPHIBIANS 179

1915. Eolodactylus aculeatus Calabresi, Monit. Zool. Ital. (Firenze) 26,

p. 238, fig. 2: Somalia.
Range. Ethiopia and British Somaliland, south through So-
malia to Kenya Colony.

Family GEKKONIDAE

Genus STENODACTYLUS Fitzinger

1826. Stenodactyliis Fitzinger, Xeue Class. Rept, pp. 13, 47. Type by

tautonomy : S. elegans Fitzinger, new name for Asmlahotes

stenodactyliis (sic) Lichtenstein.
1842. Tolarcnta Gray, Zool. Misc., p. 58. Type by monotypy: T. wilkin-

sonii Gray = Ascalabotes sthenodactylus Lichtenstein.
1874. Ceramodactylus Blanford, Ami. Mag. Nat. Hist., (4) 13, p. 454.

Type by monotypy: C. doriae Blanford.

Stenodactylus sthenodactylus sthenodactylus (Lichtenstein)
1823. Asmlahotes sthenodactylus Lichtenstein, Verz. Doiibl. Mus, Zool.
Berlin, p. 102: Egypt and Nubia, i.e. Sudan.

1823. Agame ponctue Geoffroy Saint-Hilaire, Rept., in Descr. Egypte, 1,
p. 129, pi. V, fig. 2: Egypt.

1826. Stenodactylus elegans Fitzinger (new name for Ascalabotes steno-

dactylus (sic) Lichtenstein), Neue Class. Rept., p. 47.

1827. Trapelus Savignyi Audouin, Hist. Nat. Rept., in Descr. Egypte, 1,

p. 167, and Suppl. Rept., pi. i, figs. 31—33; Egj-pt.
1829. Stenodactylus guttatus Cuvier, Regne Animal, ed. 2, 2, p. 58, pi.

iv, fig. 2 : Egypt.
1842. Tolarenta Wilkinsonii Gray, Zool. Misc., p. 59: Egypt.
1943. Stenodactylus stenodactylus savattari Seortecci, Miss. Biol. Sagan-

Omo, Zool., 1. pp. 268, 294: Elolo, Olo River, Lake Rudolf,

EtMopia.2^
Range. Syria ; Arabia ; Egypt, west to Tunisia, south to Lake
Rudolf, Kenya Colony.

23 Seortecci (loc. cit.), while stating that his two specimens from Lake Rudolf
agree m most respects with S. s. sthenodactylus, claims that they may be dif-
ferentiated by two characters. His North African material had 9-10 upper labials,
his Ethiopian geckos 13-14. The counts for twelve North African 8. s. stheno-
dactylus examined for my 1947 revision of the GEKKONIDAE, ranged from
9-15 (p. 46).

A new character used by Seortecci is the number of granules around midbody.
In his North African specimens these were 85 and 96 respectively; in his
Ethiopian geckos 105 and 106. Owing to the small and irregular size and dispo-
sition of these granules it is difficult to obtain the same number in two counts,
but our material gives the following :

$ (M.C.Z. 31395) Port Sudan, Sudan 91 or 95
9 (M.C.Z. 51659) Ein el Weibeh, Israel 98 or 100
9 (M.C.Z. 21909) Sfax, Tunisia 102 or 105

$ (M.C.Z. 5234) "Dalmatia" 119 (A.L. and B.S.)

180 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus CNEMASPIS Straucli

1887. Cnemaspis Strauch, Mem. Acad. Imp. Sci. St.-Petersbourg, (7)
35, No. 2, p. 4. Type by nionotypy: C. hoiilengeri Strauch.

1921. Paragon<itodes Noble, Amer. Mus. Novit., No. 4, p. 14. Type by
original desiguation : Gonatode.t dicTcersoni Schmidt ^= G. qnat-
tuorseriatus Sternfeld.

Cnemospis quottuorseriatas (Sternfeld)

1912. Gonatodes quattuorseriatus Sternfeld, Wiss. Ergebn. Deut. Zentral-

Afiika-Exped. 1907-1908, 4. p. 202, pi. vi, fig. 1 : Rugege Forest

and Kisenyi, Lake Kivu, Belgian Kuanda-Urundi; Uvira, Lake

Tanganyika, Belgian Congo.

1919. Gonatodes diclcersoni Schmidt, Bull. Amer. Mus. Nat. Hist., 39,

p. 436, fig. 6: Medje, Ituri District, Belgian Congo.
1950. Onemaspis bohmanni Miiller & Uthmoller, Zool. Anz., 145. p. 118:
Mto-wa-Mbu, 990 metres, at foot of Great Rift Valley about 2
km. north of Lake Manyara, Tanganyika Territory.
Range. Kenya Colony, south to central Tanganyika Territory,
west through Uganda and Belgian Ruanda-Urundi to the Belgian
Congo.

*&^

Cnemaspis airicona elgonensis Loveridge
Elgon Forest-Gecko
1936. Cnemaspis africanus elgonensis Loveridge, Proc. Zool. Soc. Lon-
don, p. 820: Above Sipi, western slopes of Mount Elgon,
Uganda.
Range. Western Kenya Colony to Mount Ruwenzori, western
Uganda.

Cnenxaspis africana africana (Werner)
Usambara Forest-Gecko
1895. Gymnodactylus africanus Werner, Verb. Zool. Bot. Ges. Wien, 45.
p. 190, pi. V, fig. 5 : Usambara Mountains, Tanganyika Terri-
tory.
Range. Central Kenya Colony, south to eastern Tanganyika
Territory.

The locality of this last gecko is no doubt erroneous, it having been received
from a dealer in February. 1884.

LOVERIDQE: E. AFRICAN REPTILES AND AMPHIBIANS 181

Genus HEMIDACTYLUS Oken

1817. Ilemidactyles Cuvier, E^gne Animal, 2, p. 47. Only included species

the "G. Tubereuleux de Daud.," i.e. Gecko tuhercidosxis Daudin

= Geclco mabouia Moreau de Jonnes.
1817. "Hemidact" von Oken, Isis, col. 1183. Assumed abbreviation for

Hemidactylus.-* No species mentioned.
182.1. HemidacUjlm Gray, Ann. Philos., 26. p. 119, "Old Continent. H.

tuberculos^is Gray. Geclco. Daud."
1827. Reviidactylus Gray, Philos. Mag., (n.s.) 2, p. .55. The only species

included is "Geclco tuberculoses Daud."

1842. Boltalia Gray, Zool. Misc., p. 58. Type by monotypy: B. sublaevi.s

Gray.

1843. Iloplopodion Fitzinger, Syst. Eept., pp. 19, 103. Type by original

designation: Hemidactylus coctaei Dumeril & Bibron.
1843. Mierodactylus Fitzinger, Syst. Eept., pp. 19, 104. Type by original

designation : Hemidactylus periivmmis Wiegmann.
1843. Onychopus Fitzinger, Syst. Eept., pp. 19,104. Type by original

designation: Hemidactylus garnotii Diimeril & Bibron.
1843. Tochyhates Fitzinger, Syst. Eept., pp. 19, 105. Type by original

designation: Geclco mabouia Moreau de Jonn6s.
1843. Pnoepus Fitzinger, Syst. Eept., pp. 19, 106. Type by original

designation : Hemidactylus javanicus Cuvier = H. frenatui

Dumeril & Bibron.
1845. Velernesia Gray, Cat. Lizards Brit. Mus., p. 156. Type by mono

typy: V. richardsonii Gray.
1845. Doryura Gray, Cat. Lizards Brit. Mus., p. 156. Type by subsequent

designation : D. bowringii Gray.
1845. Leiurus Gray (not of Hemprich & Ehrenberg), Cat. Lizards Brit.

Mus., p. 157. Type by monotypy: L. ornatus Gray = Hemidac-
tylus f. fasciattis Gray.
1845. Nubilia Gray, Cat. Lizards Brit. Mus., p. 273. Type by monotypy:

N. argentii Gray = Hemidactylus depressus Gray.

24 Oken, though only contrasting his own classlflcation with that of Cuvier,
frequently latinized the latter's vernacular names. In this instance he lists
"Gecko . . . Thecadactylus Hemidact. Platydact." The assumption that he
intended "ylus" endings, rather than "yles," is open to question.

In his description of Gecko tuberculoatis, Daudin (1802) states that It has nar-
row scales heneath the tail : this agrees with mabouia Moreau de Jennys and
excludes the confusion of some authors with Gecko gecko of Siam, for the latter
has transverse rows of small quadrangular scales beneath its tail. Furthermore,
under "G. mabuia Nob." Cuvier remarks in the 1820 edition of R6gne Animal,
that this gecko is called Mabouia des murailles in the islands, being found in all
the warmer parts of America. A footnote adds that this is the "Mabouia des
murailles" recently monographed by Moreau de Jonn6s. I am not sure why
Sherborn regarded Hemidactylus Gray, 1825, as a nomen nudum unless it Is on
account of the erroneous "Old Continent." For Gray furnishes a brief descrip-
tion, though possibly not diagnostic. The first use of the name Hemidactj/lut
approved by Sherborn is that ot Gray, 1827.

182 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1846. Hoplopodium Agassiz, Nomen. Zool., Index Univers., p. 185.

Emendation for Eoployodion Fitzinger.
1801. Eurhous Fitzinger, Sitzb. Akad. Wiss. Wien, 42. p. 400. Type b.v

monotypy : Hemidactyliis leschenaidtii Diimeril & Bibron.
18G2. Liurus Cope (not of Ehrenberg:1828), in Slack, Handbook Mus.

Acad. Nat. Sci. Philadelphia, p. 32. New name for Leiitrus Gray

(preoccupied).
1869. Teratolepis Giinther, Proc. Zool. Soc. London, p. 504. Type by

monotji^y: Hormonota fasciata Blyth.

1894. Bwnocncmis Giinther, Proc. Zool. Soc. London, p. 85. Type by

monotypy: B. modestus Giinther.
1934. Lophopholis Smith & Deraniyagala, Ceylon Journ. Sci., (B) 18,

p. 235. Type by original designation: Teratolepis scahriceps

Annandale.
1940. Aliurus Dunn & Dunn, Copeia, p. 71. New name for Liitrvs Copo

(preoccupied).

Hemidactylus olbopunctatus Loveridge

1942. Teratolepis taylori Parker (not Hemidactylus taylori Parker:

1932), Bull. Mus. Comp. Zoo!., 91. p. 33: Haud at 8° 15'N., 46°

20'E., 2100 feet, British Somalilaud.
1947. Hemidactylus alhopunctatus Loveridge (new name for Teratolepis

taylori Parker: preoccupied), Bull. Mus. Comp. Zool., 98, p. 107.
Range. British Somalilancl, south to northern Kenya Colony.

Hemidactylus isolepis Boulenger

1895. Hemidactylus isolepis Boulenger, Proc. Zool. Soc. London, p. 531,

pi. xxix, fig. 1 : Turfa Tug, south of Harar, Ethiopia.
Range. Ethiopia and British Somaliland, south through So-
malia to Kenya Colony. ,

Hemidactylus modestus (Giinther)

1894. Bunocnemis modestus Giinther, Proc. Zool. Soc. London, p. 95, pi.

viii: Ngatana, Tana River, Kenya Colony.
Range. Kenya Colon}' (known only from the type locality).

Hemidactylus tropidolepis Mocquard

1888. Hemidactylus tropidolepis Mocquard, Mem. Cent. Soc. Philom.
Paris, p. 133: Somaliland.

Range. British Somaliland, south to the Tana River, Kenya
Colony.

Hemidactylus squamulatus squamulatus Tornier*^

23 Though at one time I regarded this and the following form as races of
tropidolepis Mocquard, since collecting the latter it seems advisable to treat the
much larger squamulatus as distinct.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 183

189(5. Eeniidactylus squamulatus Tornier, Die Kriechthiere Deutsch-Ost
Afrikas, p. 10: Kakoma, Uguuda, Tanganyika Territory.

1896. Hemidaciylus bocagci Tornier (not of Boulenger), Die Kriech-

thiere Deutsch-Oat-Afrikas, p. 12: Dalalani, Lake Natron, Tan-
ganyika Territory.

1897. Remidactylus tverneri Tornier (new name for H. hocagei Tornier:

preoccupied), Arch. Naturg., 63, Abt. 1, p. 63.
1902. Hemidaciylus Tornieri Moequard (new name for R. hocagei

Tornier: preoccupied), Bull. Mus. Hist Nat. (Paris), 8. p. 407.
1923. Hemidactyhts Alluaudi Angel, Bull. Mus. Hist. Nat. (Paris), 29,

p. 490: Bura, Teita, Kenya Colony.
Range. Northeastern Kenya Colony, south to central Tan-
ganyika Territory.

Hemidaciylus squamulatus barbouri Loveridge

1942. Uemidactyhis tropidolepis harhouri Loveridge, Bull. Mus. Comp.
Zool., 91. p. 320, fig. — : Changamwe, Kenya Colony.

Range. Coastal Kenya Colony from Malindi, south to Tanga.
Tanganyika Territory.

Hemidaciylus frenatus Dumeril & Bibron

1830. Hemidactylus frenatus "Schlegel" Dumeril & Bibron, Erpe1.
Gen., 3. p. 366: Java (restricted); South Africa, etc.

1843. Hemidactylus (Fhoepus) javaniciis "Cuvier" Fitzinger, Syst.
Eept., p. 106: Java (restricted).

1843. Hemidactylus (Fhoepus) Bojeri Fitzinger, Syst. Eept., p. 106:
Cape of Good Hope; Madagascar; Mauritius.

184.5. Hemidactylus vittatus Gray, Cat. Lizards Brit. Mus., p. 15-5:
Borneo.

1854. ^.Hemidactylus punctatus Jerdon, Journ. Asiatic Soc. Bengal, 22, p.
467 (type lost) : Tellicherry, Malabar, India.

1860. Hemidactylus inornatus Hallowell, Proe. Acad. Nat. Sci. Phila-
delphia, p. 492: Eiu Kiu Islands.

1860. Hemidactylus pumilus Hallowell, Proe. Acad. Nat. Sci. Phila-
delphia, p. 502 : Hongkong.

1864. Geclco chaus Tytler, Journ. Asiatic Soe. Bengal, (2) 33, p. 547:
Eangoon, Burma (restricted).

1864. Geclco caracal Tytler, Journ. Asiatic Soc. Bengal, (2) 33, p. 547:
Rangoon, Burma.

1868. Hemidactylus longiceps Cope, Proe. Acad. Nat. Sci. Philadelphia,
p. 320: Manila, Philippine Islands.

1868. Hemidactylus hexaspis Cope, Proe. Acad. Nat. Sci. Philadelphia,
p. 320 : Madagascar.

184 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1915. Hemidactylus fragilis Calabresi, Monit. Zool. Ital. (Firenze),
26. p. 236, fig. 1 : Bur Meldac, Somalia.

Range. Somaliland ; Lamu Island ; Seychelles ; Mauritius ;
Madagascar; St. Helena and other islands of the Indian and
Pacific Oceans ; Ceylon ; southern India ; Malay Peninsula ; Indo-
China; China; Korea (This wide distribution may be attributed
to human agency).

Hemidactylus puccionii Calabresi

1927. Hemidactylus puccionii Calabresi, Atti. Soc. Ital. Sci. Nat.

(Milano), 86. pp. 23, 39, pi. i, figs. 3-3b: Obbia to Durgale,

Somalia.
1936. Hemidactylus parJceri Loveridge, Proc. Biol. Soc. Washington, 49.

p. 59: Zanzibar Island (? introduced). •

Range. Somalia and Zanzibar Island ( ? introduced ) .

Hemidactylus citernii Boulenger

1912. Hemidactylus citernii Boulenger, Ann, Mus. Civ. Stor. Nat. Genova,
(3) 5. p. 329: Rahanuin country, Somalia.

Range. British Somaliland, south through Somalia to Kenya
Colony ( ? introduced. Erroneously reported from Tanganyika
and Zanzibar).

Hemidactylus brookii angulatus Hallowell

1852. Hemidactylus angidatus Hallowell, Proe. Acad. Nat. Sci. Phila-
delphia, p. 63, fig. — : "West coast of Africa," i.e. Gabon,
French Congo.
1868. Hemidactylus guineensis Peters, Monatsb. Akad. Wiss. Berlin, p.

640: "Ada Foah, Guinea," i.e. Adafer, Mauritania.
1870. Hemidactylus affinis Steindachner, Sitzb. Akad. Wiss. Wien, 62.

Abt. 1, p. 328 : Dagana and Goree, Senegal.
1885. Hemidactylus stellatus Boulenger, Cat. Lizards Brit. Mus., 1. p.

130, pi. xii, fig. 1: Gambia and West Africa.
1893. Hemidactylus Bayonii Bocage, Jorn. Sci. Lisboa, (2) 3. p. 116:

Dondo, Cuanza River, Angola.
1897. Hemidactylus hrooTcii var. Togoensis Werner, Verh. Zool. Bot. Ges.

Wien, 47. p. 397: Atakpanie, Togo.
Range. Sudan and Uganda, south to Tanganyika Territory,
west through Belgian Congo to Angola and French Congo,
northwe-st to Senegal and the Cape Verde Islands. (Its distribu-
tion is due in part to human agency. Another race occurs in the
West Indies and South America; two in Asia.)

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 185

Hemidactylus turcicus macropholis Boulenger

1896. Eeviidactylus macropholis Boulenger, Ann. Mus. Civ. Stor. Nat.
Genova, (2) 17, p. 7, pi. i, figs. 2-2c: Dolo, Somalia.

Range. Ethiopia and British Somalilaud, south through Som-
alia to Kenya Colony. (Another race occurs in Arabia, as does
the typical form whose wide distribution in four continents is
in part due to human agency.)

Hemidactylus ruspolii Boulenger

1890. Hemidactylus ruspolii Boulenger, Ann. Mus. Civ. Stor. Nat.
Genova, (2) 17, p. 6, pi. i, figs. 1-lc: Bardera; Dolo; Lugh;
Magala and Umberto Island, Somalia.
1907. Hemidactylus erlangeri Steindachner, Ann. Akad. Wiss. Wieu, 44.

p. 355 ; Ethiopia.
Range. Ethiopia and British Somaliland, south to northern
Kenya Colony.

Hemidactylus tanganicus Love ridge

1929. Hemidactylus tanganicus Loveridge, U. S. Nat. Mus. Bull., 151,
p. 42, pi. i: Dutumi, near Kisaki, Tanganyika Territory.

Range. Tanganyika Territory (known only from the large
holotype).

Hemidactylus mobouia (Jonnes) Common House-Gecko

1802. Geclco tuherculosus Daudin, Hist. Nat. Eept., 4. p. 158, footnote:

No locality.
1818. GecTco Mabouia Moreau de Jonnes, Bull. Soc. Philom. Paris, p. 138:

Antilles and adjacent mainland.
1825. GecTco aculeatus Spix, Animalia Nova Species Novae Lacerta., p.

16, pi. xviii, fig. 3: Rio de Janeiro, Brazil.
1825. GecTco cruciger Spix, Animalia Nova Species Novae Lacerta., p. 16:

Bahia, Brazil.
1825. GeTclco incanescens Wied, Beitr. Naturg. Brasilien, 1, p. 101: Brazil.
1825. GeTcTco armatus Wied, Beitr. Naturg. Brasilien, 1, p. 104: Brazil.
1854. '^Hemidactylus platycepTialus Peters, Monatsb. Akad. Wiss. Berlin,

p. 615: Coast north of Mozambique; Johanna Island, Comoro

Islands.
1867. Hemidactylus SaTcalava Grandidier, Revue Mag. Zool., (2) 19. p.

233 : Tulear, Madagascar.
1879. Hemidactylus frenatus var. calabaricus Boettger, Ber. Offenbach.

Ver. Naturk., 17-18, p. 1: Old Calabar, Nigeria.
1893. Hemidactylus benguellensis Bocage, Jorn. Sci. Lisboa, (2) 3, p.

115: Cahata, Benguela, Angola.

186 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1932. Hcmidactylus tasmani Hewitt, Ami. Natal Mus., 7, p. 120 : Drief on-
tein, near Gwelo, Soutliern Ehodesia.

Range. Possiljly Eritrea and Ethiopia; c'ertaiiil,y Somalia
south throuoh Kenva (Jolonv; Tano'anvika Territory; Pemba,
Zanzibar and Mafia LslancLs to Zuluhmd ; north^Yest through
Transvaal and Bechnaualand to Angola ; north and northwest to
Liberia. (In West Africa, however, the distribution is spotty,
presumably due to introduction at ports.) West Indies. Mexico.
South America. Madagascar.

Hemidactylus mercatorius Gray Coconut-Palm Gecko

1842. Hemulact ijlns mercatorius Gray, Zool. Misc., p. 58: Madagascar.
1909. Uemidactylus gardin.cri Boiileiiger, Trans. Linn. Soc. London, (2)

12. p. 296, pi. xi, fig. 4: Farquhar Lsland, Seychelle Islands.
1928. II emida ct yhis pcrsimilis Barbour & Loveridge, Mem. Mus. Conip.
Zool., 50, p. 140, pi. iv, figs. 1, 3: Dar es Salaam, Tanganyika
Territory.
19.')o. Eemidactylus n'tmdanus Loveridge, Proc. Biol. 8oc. Washington,

43, p. 60 : Kitau, Manda Island, Kenya Colony.
Range. Coastal Kenya Colony and Tanganyika Territory ; in-
cluding Pemija Island (and almost certainly Zanzibar and Mafia
Islands), the Sej^chelle and Aldabra Islands, Madagascar and
Mauritius; south to Mozambique, inland to Nyasaland.

Genus LYGODACTYLUS Gray

1864. Lygodactylus Gray, Proc. Zool. Soc. London, p. 59. Type by
monotypy: L. strigatus Gray = Hemidactylus capensis A. Smith.

1880. Scalabotes Peters, Monatsb. Akad. Wiss. Berlin, p. 795. Type by
monotypy : 5. thomensis Peters.

1883. Microscalahotes Boulenger, Ann. Mag. Nat. Hist., (5) 11, p. 174.
Type by monotypy: M. cowani Boulenger.

Lygodactylus somalicus Loveridge

1935. Lygodactylus somalicus somalicus Loveridge, Proc. Biol. Soc.
Washington, 48, p. 196: Bar Madobe, Nogal Valley, British
Somaliland.
1935. Lygodactylus somalicus annectens Loveridge, Proc. Biol. Soc. Wash-
ington, 48, p. 197: Buran District, British Somaliland.
Range. British Somaliland, south through Somalia to Kenya
Colony.

Lygodactylus schefileri Sternfeld

1912. Lygodactylus fischeri scheffleri, Sterireld, Wiss. Ergebn. Deut.

LOVERIDGE: E. AFRICAN * REPTILES AND AMPHIBIANS 187

Zentral-Afrika-Exped. 1907-1908, A. p. 206: Kibwezi, Kenya
Colony.
Range. Kenya Colony.

Lygodactylus conradti jMatschie

1892. Lygodactylus conradti Matschie, Sitzb. Ges. Naturf. Freunde Ber-
lin, p. 109: Dercma, Usambara Mountains, Tanganyika Terri-
toiy.

Range. Kenya Colony and Tanganj^ka Territory.

Lygodactylus angolensis Boeage

1896. Lygodactylus angolensis Boeage, Jorn. Sci. Lisboa, (2) 4. p. 110:

Hanha, Benguela. Angola.
1926. Lygodactylus stcvc7isoni Hewitt, Ann. Natal Mus., 5. p. 445, pi.

x.\v, figs. 3-4: Khami Ruins, Bulawayo, Southern Rhodesia.
1933. Lygodactylus laurae Schmidt, Ann. Carnegie Mus., 22, p. 4: Chitau,

Bihe District, Angola.
Range. Tanganyika Territory, south to Southern Rhodesia,
west througli Beehuanaland to Angola and the Belgian Congo.

Lygodactylus capensis (>Smith) Cape Dwarf-Gecko

1849. Ilcmidactylus capensis A. Smith, Illus. Zool. S. Africa, Rept., pi.

Ixxv, fig. 3 : Kaffirland and districts north of Cape Province.
18G4. Lygodactylus strigatu-s Gray, Proc. Zool. Soc. London, p. 59:

"Southeastern Africa." i.e. Zambezi region.
1932. Lygodactylus bradfieldi Hewitt, February, Ann. Natal Mus., 7,
p. 126, pi. vi, fig. 10: Quickborn Farm, near Okahandja, South-
west Africa.
1932. Lygodactylus capensis ngarnieyisis FitzSimons, October, Ann.
Transvaal Mus., 15, p. 35: Mabeleapudi, Ngamiland, Beehuana-
land Protectorate.
Range. Kenya Colony and Tanganyika Territory, south to
Natal, west through northern Cape Province to Southwest Africa
and Angola.

Lygodactylus grotei grotei Sternfeld

1911. Lygodactylus Grotei Sternfeld, Sitzb. Ges. Naturf. Freunde Ber-
lin, p. 245 : Mikindani, etc., Tanganyika Territory.
1920. Lygodactylus capensis mossamhica Loveridge, Proc. Zool. Soc.

London, p. 135: Lumbo, Mozambique.
Range. Tanganyika Territory and Mafia island, south to Mo-
zambique.

Lygodactylus grotei pakenhami Loveridge
Pemba Island Dwarf-Gecko

1 88 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1941. Lygodactylus grotei paTcenhami Loveridge, Proe. Biol. Soc. Wash-
ington, 54. p. 176: Wete, Pemba Island,
Range. Pemba Island.

Lygodactylus angularis angularis Giinther

1893. Lygodactylus angularis Giinther, Proe. Zool. Soc. London, 1892, p.
555, pi. xxxiii, figs. 1-3: Shire Highlands, Nyasaland.

Range. Tanganyika Territory and Nyasaland, west to North-
ern Rhodesia.

Lygodactylus picturatus gutturalis (Bocage)

1873. Hemidactylns gutturalis Bocage, Jom. Sci. Lisboa, 4. p. 211:
Bissau, Portuguese Guinea.

Range. Uganda and Tanganyika Territory (Ujiji only), west
to French Congo. Portuguese Guinea.

Lygodactylus picturatus keniensis Parker

]93(). Lygodactylus picturatiLS heniensis Parker, Ann. Mag. Nat. Hist.,

(10) 18, p. 602: Lodwar, Lake Rudolf, Kenya Colony.
Range. Kenya Colony (north of the Uaso Nyiro).

Lygodactylus picturatus uikerewensis Loveridge

1935. Lygodactylus picturatus uTcerewensis Loveridge, Proe. Biol. Soc.
Washington, 48. p. 199: Ukerewe Island, Lake Victoria, Tan-
ganyika Territory.
Range. Kenya Colony and Tanganyika Territory (in vicinity
of Lake Victoria).

Lygodactylus picturatus mombasicus Loveridge

1935. Lygodactylus picturatus mombasicus Loveridge, Proe. Biol. Soc.
Washington, 48. p. 198: Kilindini, Mombasa Island, Kenya
Colony.
Range. Coastal Kenya Colony (meets with typical form at
both Mombasa and Tanga), south to Tanga, Tanganyika Terri-
tory.

Lygodactylus picturatus picturatus (Peters)
Yellow-headed Dwarf-Gecko

1868. Hemidactylus variegatus Peters (not of Dumeril & Bibron:1836),
Monatsb. Akad. Wiss. Berlin, p. 449: "Zanzibar Coast," i.e.
Tanganyika Territory.

1870. Hemidactylus picturatus Peters (new name for variegatus Peters:
preoccupied), Monatsb. Akad. Wiss. Berlin, p. 115.

1896. Lygodactylus picturatus vars. griseus, septcmlineatus and quin-

LOVEBIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 189

quelineatus Tornier (nomina nuda), Die Kriechthiere Deutsch-
Ost-Afrikas, p. 15: No tj'pes or type localities designated.
1928. Lygodactylus Duvnni Loveridge, Proc. U. S. Nat. Mus., 72, Art.

24, p. 1, pi. i: Saranda, Ugogo, Tanganyika Territory.
Range. (Somalia ? or) Kenya Colony (coastal belt from
Mombasa) south to Tanganyika Territory and Mozambique
(chiefly coast, rivers and railways) ; Zanzibar and Mafia Islands.
Northern Rhodesia and neighboring parts of the Belgian Congo
(its present distribution has been complicated by transportation
as eggs or adults through human agency).

Lygodactylus picturatus williamsi Loveridge
Turquoise-Blue Dwarf-Gecko
1952. Lygodactylus picturatus tvilliams-i-^ Loveridge, Jouru. E. Africa
Nat. Hist. Soc, vol. "xx," p. "446," = 21. p. 39: Kimboza
Forest, 1000 feet. Eastern Province, Tanganyika Territory.
Range. Tanganyika Territory (known only from the holo-
type).

Genus PHYLLODACTYLUS Gray=^

1828. Phyllodactylus Gray, Spicil. ZooL, p. 3. Type by monotypy: P.

pv.lcher Gray.
1843. Eideptes Fitzinger, Syst. Rept., pp. 18, 95. Type by original

designation : Phyllodactylus europaeus Gene.
1843. Discodactylus Fitzinger, Syst. Rept., pp. 18, 95. Type by original

designation: Pliyllodactyhts tuberculosvs Wiegmaiin.
1845. Gerrliopygus Gray, Cat. Lizards Brit. Mus., p. 150. Type by tan-

tonomy: Diplodactylus gerrhopygus Wiegmaun.
1879. Paroedura Giinther, Ann. Mag. Nat. Hist., (5) 3. p. 218. Type by

monotypy: P. sanctijohamiis Giinther.

Phyllodactylus wolterstorffi (Tornier) ^^

1900. Diplodactylus wolterstor-jfl Tornier, Zool. Jalirb., Syst., 13, p. 584,
fig. A: Tanga, Tanganyika Territory.

26 Altered by the then editor to Williamsi in some printings, to villiamsi iu
others ; both volume number and pagination were in error, so corrected in a subse-
quent issue by the new editor.

2" Here used in the new sense proposed by Garth Underwood (1954, Proc. Zool.
Soc. London, 124, p. 472) involving the transfer to Phyllodactylus of those
species with a Gekko-type pupil formerly referred to Diplodactylus, a genus now
restricted to Australia.

28 Through the kindness of Dr. Heinz Wermuth, I recently had the opportunity
of examining the type and a paratype (now M.C.Z. 54700). They reveal some
minor defects in Tornier's description, but though there is little to differentiate
them from P. inexpectatus (Stejneger )of the Seychelles, direct comparison with
a specimen of the latter (M.C.Z. 49198) shows that the two are distinct.

1!*0 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

Range. Tanganyika Territory (known only from tlie four
types).

Genus EBENAVIA Boettger

1S78. Ebenavia Boettger, Abliaud. Senekenberg. Naturf. Ges., II, p. i]7t>.
Type by monotypy: E. inunguis Boettger.
Ebenavia sp.'^

Range. Pemba Island.

(lenus PHELSUMA Gray

182rj. riiclsiiina Gray, Ann. Philos., (2) 10. p. 199. Type by monotypy:
•'P. crepidianus," i.e. GecJco cepedianus Merrem.

1830. Anoplopus Wagler, Nat. Syst. Amphib., p. 142. Type by designa-
tion of Fitzinger :1843 : GecJ:o ccpedianiiJi Merrem.

Phelsuma dubia dubia (Boettger)

1881. Padiydartylu,s duhius Boettger, Zool. Anz., 4, p. 4G : Madagascar.

Range. Coastal Tanganyika Territory; Zanzibar Island ; Com-
oro Islands; northwest Madagascar.

Phelsuma madagascariensis parkeri Loveridge

1911. Phelsuma madagascariensis parheri Loveridge, Proe. Biol. Sdc

Washington, 54. p. 17.5: Kinowe, Pemlia Island.
Range. Pemba Island.

Genus HOMOPHOLIS Boulenger

188.^). Eoiiiopholis Boulenger, Cat. Lizards Brit. Mus., 1, p. 191. Type
l)y monotypy: Gecko walhci-gi (sic) A. Smith.

-9 Admission of this genus to the East African list rests solely on some geckos
iiewlv hatched from esKs, measuring 5x6 mm., found at Cholvocho and Kinanga-
juu, Pemba Island, by R. H. Pakeuham (1947, Ann. Mag. Nat. Hist., (11) 14,
V- 135). The assignment to Ehcnaiia is unquestionably correct; it was made by
n. W. Parker. At my request he very kindly re-e.xamined them under eery high
magnification supijlying me witli a sketch and the substance of the following :

Scales of head nuilticarinate, uniform, their size corresponding to the larger
liody granules : the latter have some indications of a keel, all the dorsal granules
appearing to be sul)iml)ricate with a surface sculpturing of minute papillae ;
among tliem are irregular rows of large rounded tubercles ; no enlarged chin
sliields ; no claws. Color above, pale straw, with or without a few, very
faint, longitudinal lines that converge upon the iiase of tail : the dorsal color
sharply demarcated from that on the sides of head, neck and flanks, this is dark
sepia passing gradually into the light sepia of the undersurface : tail above dis-
plays some paired spots anteriorly, while posteriorly it is banded with darker and
ligliter. This is, of course, hatchling coloration and probably differs from that
of the adult. Capture of some adults on I'emba is necessary to settle the point
as to whether the species is distinct, or whether inunguis Boettger has been
introduced fruni Xosy IW-. Rladagascar. and become established on Pemba.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 191

1890. Platjpholis Boulenger, Proe. Zool. Soc. London, p. 80, pi. vii,
fig. 2. Type by monotypy: P. fasciata Boulenger.

Homopholis fasciata lascicrta (Boulenger)

1890. Platypholis fasciata Boulenger, Proc. Zool. Soc. London, pp. 77.
81, pi. viii, fig. 2: Mombasa, Kenya Colony.

Range. Nortlieni Kenya Colony, south to Tanganyika Terri-
tory.

Genus PACHYDACTYLUS Wiegmann

1834. Pachy dactyl us Wiegmann, Herp. Mexicana, p. 19. Type by mono-
typy: P. hcrgii Wiegmann = Laxierta geitje Sparrman.

1843. Colobopus Fitziuger, Syst. Eept., p. 19. Type by original designa-
tion: Anoplopus inunguis Wagler := Gecko inimgnis Cuvier,
1817 = Lacerta geitje Sparrman, 1778.

1845. Cantinia Gray, Cat. Lizards Brit. Mus., pp. 167, 168. Type by
monotypy: Pachydactylus elegans Gray = Tarentola capensis
A. Smith.

1864. Eomodactylus Gray (not Fitzinger :1843), Proc. Zool. Soc. London,
p. 59. Type by monotypy: H. turneri Gray.

1894. Elasmodactylus Boulenger, Proe. Zool. Soc. London, p. 724. Tj'pe
by monotypy: E. tubereulosiis Boulenger.

Pachydactylus bibronii turneri (Gray)

1864. Eomodactylus turneri Gray, Proc. Zool. Soc. Loudon, p. 59, pi. ix,

fig. 2 : Tete, Mozambique.
1910. Pachydactylus bibronii var. st.ellatus Werner, Denks. Med.-Nat.

Ges. Jena, 16, p. 309: Great Namaqualand, Southwest Africa.
Range. Belgian Ruanda-Urundi and Tanganyika Territory,
south to Mozambique, west through Nyasaland ; the Rhodesias ;
Transvaal ; Bechuanaland and Orange Free State to Little Nama-
qualand and adjacent Cape Province, north through Southwest
Africa to southern Angola.

PachydactYlus tetensis Loveridge

19 '12. Pachydactylus tetensis Loveridge, Bull. Mus. Comp. Zool., 110,
p. 17.5, pi. V, fig. 3: MAvanza Eocks, Kasumbadedza, near Tete.
Mozambique.
Range. Near Liwale, Tanganyika Territory, south to Mozam-
bique (on south bank of the Zambezi).

Pachydactylus tuberculosus (Boulenger)

1894. Elasmodactylus tuberculosus Boulenger, Proc. Zool. Soc. London.

192 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

pp. 723, 727, pi. xlvii, fig. 2: Lower Congo.
1896. Pachydactylus boulengeri Tornier, Die Kriechthiere Deutsch-Ost-

Af rikas, p. 26, pi. ii, fig. 1 : Tabora and Kakoma, Tanganyika

Territory.
1913. Elasmodactylus triedrus Boulenger, Eevue Zool. Afr., 3, p. 104,

figs. — : Kikondja, Katanga, Belgian Congo.
Range. Tanga, Tanganyika Territory, southwest to Nyamkolo,
Northern Rhodesia ; west to Lower Congo River ; Belgian Congo.

Family AGAMIDAE
Genus AGAMA Daudin'°

1802. Agama Daudin (part). Hist. Nat. Eept., 3. pp. 333, 356. Type by
subsequent designation: A. colonorum Daudin (part) = Lacerta
agama Linnaeus.

1817. Trapelus Cuvier, Eegne Animal, 2, p. 35. Type by monotypy:
Agame variahile Geoffrey = Agama m.utabilis Merrem.

1826. Tapaya Fitzinger, Neue Class. Kept., pp. 17, 49. Type by subse-
quent designation : Agama orbicularis Cuvier = Lacerta hispida
Linnaeus.

1833. Cyclosaurus Wagler, Isis von Oken, col. 894. Type by original
designation : Agama orbicularis Daudin = Lacerta hispida
Linnaeus.

1843. Phrynopsis Fitzinger, Syst. Kept., pp. 17, 79. Type by original
designation: Agama atra Daudin.

1843. Podorrhoa Fitzinger, Syst. Kept., pp. 18, 80. Type by original
designation: Agama colonorum Daudin = Lacerta agama Lin-
naeus.

1843. Pseudotrapelus Fitzinger, Syst. Kept., pp. 18, 81. Type by original
designation: AgamM sinaita Heyden.

1843. Planodes Fitzinger, Syst. Kept., pp. 18, 81. Type by original
designation: Agama agilis Olivier.

1843. Trapeloidis Fitzinger, Syst. Kept., pp. 18, 81. Type by original
designation: Lacerta sanguinolenta Pallas.

1843. Psammorrhoa Fitzmger, Syst. Kept., pp. 18, 81. Type by original
designation: Agama acvleata Merrem.

1843. Eremioplanis Fitzinger, Syst. Kept., pp. 18, 82. Type by original

30 Elimination of Stellio Laurenti (part), 1768, Syn. Kept., p. 56, was achieved
by subsequent designation of its type as iS. saxatilis I>aurenti by Stejneser, in
M. A. Smith, 1933, Journ. Bombay Nat. Hist. Soc. for 1932, 35, p. 6l9. S.
saxatilis, being based on Seba, 1735, Locup. Rerum Nat. Thesaurus, 2, pi. Ixxix,
fig. 4, is declared unrecognizal)le. The genus SteUio has been used in other
senses by Schneider (1792) ; Latreille (1802) ; Wagler (1830) and others.

Intentionally omitted from the synonymy are obvious misspellings such as
Trnpetua Oken (1817) ; Tapelus Gray (1825) : Cyoclosaurus Wagler (1833).

IX)VERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 193

designation: Trapelus aegypiius Cuvier = Agavia mutabilis

Merrem.
1843. Acanihocercus Fitzinger, Syst. Rept., pp. 18, 84. Type by original

designation: Stellio cyanogaster Riippell.
184.3. Laudalcia Gray, Cat. Lizards Brit. Mns., p. 254. Type by mono

typy: Agama tuberculaia Gray.
1845. Isodactylus Gray, Cat. Lizards Brit. Mus., p. 2J9. Type by niono-

typy: Agama sinaita Heyden.
1854. Plocederma Blyth, Journ. Asiatic Soc. Bengal, 23. p. 738. Type

by monotypy: Laudalcia {Plocederma) melanura Blyth.
18r)6. Brachysaura Blyth, Journ. Asiatic Soc. Bengal, 25. p. 448. Type

by monotypy: B. ornata Blyth = Agama minor Hardwicke &

Gray.
1860. Barycephalus Giinther, Proc. Zool. Soc. London, p. 150. Type by

monotypy: B. syTcesii Giinther = Agama Uibercidata Gray.
1895. Xenagama Boulenger, Proc. Zool. Soc. London, p. 534. Type by

original designation: Uromastix batilliferus Vaillant.

Agama ruppelli occidentalis Parker ^^
Western Arboreal-Agama
1932. Agama rueppelli occidentalis Parker, Journ. Linn. Soc. London,
Zool., 38. p. 225 : near mouth of Kaliokwell Eiver, Lake Eudolf,
Kenya Colony.
Range. Southern Ethiopia to northern Kenya Colony.

Agama ruppelli septentrionalis Parker
Southern Arboreal-Agama
1932. Agama rueppelli septentrion-alis Parker, Journ. Linn. Soc. London.
Zool., 38, p. 225: Mount Nyero (as Njiro) ; Madago's village;
\'oi and Mbunyi, Kenj-a Colony.
Range. Central and southern Kenya Colony.

Agama hispida armata Peters^- Peters' Spiny Agama

1854. Agama armata Peters, Monatsb. Akad. Wiss. Berlin, p. 616: Rios

de Sena ; Tete, Mozambique.
Range. Tanganyika Territory, south through Mozambique;

N'yasaland and the Rhodesias to Natal.

Agama mossambica mossambica Peters ^Mozambique Agama
1854. Agama mossambica Peters, Monatsb. Akad. Wiss. Berlin, p. 616:
Coast of Mozambique.

31 A. vaillanti Boulenger is a synonym of A. r. ruppelli Vaillant of Ethiopia
and the Somalilands. Keuya records of vnillaiiti should be referred to one or
other of Parker's races.

32 Early East African records of hinpidn or hispida distantl Boulenger slioulil
be referred to this race, the other two races being South African.

194 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1874. Agama carmiventris Peters, Monatsb. Akad. Wiss. Berlin, p. 150:

Zanzibar Coast; i.e. Tanganyika Territory.
Range. Tanganyika Territory, south through northern Mo-
zambique and Nyasaland to Northern Rhodesia."^

Agama mossambica montana Barbour & Loveridge^^
Montane lloek-Agama
1928. Agama mossambica montana Barbour & Loveridge, Mem. Mus.
Comp. Zool., 50, p. 147: below Bagilo, Uluguru Mountains,
Tanganyika Territory.
Range. Usambara and Uluguru Mountains, Tanganyika Ter-
ritory .

Agama agama agama (Linnaeus) Oonnnon Rock-Agama

17.18. Laccrta agama Linnaeus (part), Syst. Nat., ed. 10, 1, p. 207:
"America" (error).

1802. Agama colonorum Daudin, Hist. Nat. Kept., 3. p. 3o6: " Amerique
meridionale, " etc. (error).

1831. Agama Occipitalis Gray, in Griffith, Animal Kingdom, 9. Synopsis,
p. 56 : Africa.

1877. Agama colonorum var. congica Peters, Monatsb. Akad. Wiss. Ber-
lin, p. 612 : Chinchoxo, Cabinda.

1877. Agama picticaiula Peters (? part^^) Monatsb. Akad. Wiss. Berlin,
p. 612: "Ada Foah, " i.e. ? Adafer, Mauretania, French West
Africa; Accra, Gold Coast; also Cameroon.

Range. Southern Sudan and Uganda, west to Nigeria, south
lo Angola.

Agama agama lionotus Boulenger Kenya Rock-Agama

1896. Agama lionotus Boulenger, Proc. Zool. Soc. London, p. 214, pi.

viii : southeast of Lake Eudolf , Kenya Colony.
Range. Eastern Uganda (Suk), southeast to Voi, Kenya Col-
ony.

Agama agama usombarae Barbour & Loveridge
Usambara Rock-Agama
1928. Agama colonorum lu^ambarae Barbour & Loveridge, Mem. Mus.

33 The two young aganias from Athi Plains, Konya Colony, referred to mossaiit-
bica by Moccinard (l'J02, Bull. Mus. Hist. Nat. (Paris), 8, p. 405) are possibly
A. a. lionotus, a species that occurs there (M.C.Z. 44260-1).

34 I have long thought that montana is a full species but, pending a thorough
revision of the entire genus, consider it is best left as described. A. »i. mossam-
hica occurs on the lower slopes of the Uluguru range.

35 While Cameroon specimens are unquestionably A. a. agama, our Gold Coasi
material seems nearer to the western race A. a. africana Hallowell. The question
cannot be settled until a series from Ada Foah has been studied, being an area
where the races meet.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 195

Conip. Zool, 50, p. 150, pi. ii, fig. 1: Soni, Usambara Mountains,
Tanganyika Territory.
Range. T^sambara Mountains, Tanganyika Territory.

Agama agama elgonis Lonnberg Elgon Roek-Agama

1922. Agama elgonis Lonnberg, Arkiv. Zool., 14, No. 12, p. 2: Mount

Elgon, Kenya Colony.
1932. Agama agama turuensis Loveridge, Bull. Mus. Conip. Zool., 72,
p. 376: Unyanganyi, Turu, Tanganyika Territory.

Range. Uganda and Kenya slopes of Mount Elgon, south to
Usandawi in central Tanganyika Territory.

Agama agama dodomae Loveridge Dodoma Rock- Agama

1923. Agama lionotus var. dodomae Loveridge, Proc. Zool. Soc. London,

p. 944: Dodoma, Ugogo, Tanganyika Territory.
Range. Central to southwestern Tanganyika Territory.

Agama agama ufipae Loveridge IJfipa Roek-Agama

1932. Agama agama ufipae Loveridge, Bull. Mus. Conip. Zool., 72, p. 377:

near Kipili, Ufipa, Tanganyika Territory.
Range. Western Tanganyika Territory (Lake Tanganyika).

Agama planiceps mwanzae Loveridge Mwanza Roek-Agama
1923. Agama lionotus var. mwanzae Loveridge, Proc. Zool. Soc. London,

p. 945 : Slianwa, Mwanza, Tanganyika Territory.
Range. Northwestern Tanganyika Territory.

Agama planiceps caudospina Meek Elmenteita Roek-Agama
1910. Agamn caudospinn Meek, Field Mus. Nat. Hist., Zool. Series, 7,

p. 407: Lake Elmenteita, Kenya Colony.
1935. Agama agama Icaimosae Loveridge, Bull. Mus. Comp. Zool., 79. p.

10: near Kaimosi, Kakamega, Kenya Colony.
Range. Kenya Colony.

Agama annectens Blanford Eritrean Roek-Agama

1870. Agama annectens Blanford, Zool. Abyssinia, p. 446, fig. — : "Soo-

roo ' ' i.e. Suru Pass, Eritrea.
Range. Eritrea, Ethiopia and the Somalilands, south to north-
ern Kenya Colony.

Agama cyanogaster (Riippell) Black-necked Arboreal-Agama
1835. Stellio cyanogaster Riippell, Neue Wirbelth. Fauna Abyssinica,

Amphib., p. 10, pi. v: Massaua, Eritrea.
1849. Agama atricollis A. Smith, Illus. Zool. S. Africa, Eept., App., p.
14: Xatal, South Africa.

196 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1851. Stellio Capensis A. Dumeril, Cat. Method. Coll. Kept. Mus. Paria,

p. 106: "Cape of Good Hope," i.e. South Africa.
1866. Stellio nigricollis Bocage, Jorn. Sci. Lisboa, 1, p. 43: Duque de

Braganca, Huila Plateau, Mossamedes, Angola.
1894. Agama gregorii Giinther, Proc. Zool. Soe. London, p. 86: Mkon-

umbi, near Lamu, Kenya Colony.
Range. Eritrea, south through Ethiopia; Uganda; Kenya
Colony and Tanganyiiva Territory (apparently absent from the
coastal islands of Pemba, Zanzibar and Mafia ! ) to Natal, north-
west through Bechuanaland to Ovamboland, Southwest Africa;
Angola and Belgian Congo.

Family CHAMAELEONIDAE^'^
Genus CHAMAELEO Laurenti

1763. CJiamaeleon Gronovius,^'^ Zoophyl. Gronov. . . ., p. 12.

1768. Chamaeleo Laurenti, Syn. Rept., p. 45. Type by subsequent desig-
nation: C. parisiensiiim Laurenti = Lacerta chamaeleon Lin-
naeus.

1839. Diceros Swainson, Nat. Hist. Fishes, Amph. Rept., 2, pp. 347, 369.
TjT)e by monotypy: D. bifurcatus Swainson.

1839. Trioceros Swainson, Nat. Hist. Fishes, Amph. Rept., 2, pp. 347,
369. Type by monotypy : T. grayii Swainson.

1843. Trioeras Fitzinger, Syst. Rept., pp. 15, 42. Type by original
designation : Chamaeleo oweni Gray.

1843. Furcifer Fitzinger, Syst. Rept., pp. 15, 42. Type by original
designation: Chamaeleon Mjidus Brongniart.

1843. Bradypodion Fitzinger, Syst. Rept., pp. 15, 43. Type by original
designation: Chamaeleon pumilus Latreille = Lacerta pumUa
Gmelin.

1846. Bradypodium Agassiz, Nomen. ZooL, Index Univers., p. 52. Emen-
dation for Bradypodion Fitzinger.

186.3. Apola Gray, Proc. Zool. Soc. London for 1864, pp. 467, 473. Type
by monotypy : Chamaeleo lateralis Gray.

36 CHAMAELEONTIDAE, derived from CHAMAELEONTES Fitzinger (1843:
41) is rejected. This family was for long referred to a separate suborder
(RHIPTOGLOSSA) on account of the peculiar projectile tongue possessed b.v all
its members. In recent years, however, this separation has been considered of
less significance than the obvious relationship to the AGAMIDAE, from which
stock chameleons appear to have been derived.

37 A worlj speciflcall.v rejected from tasonomic purposes in rule 89 of the
Internat. Comm. Zool. Nomencl. — cf. 1926, Proc. Biol. Soc. Washington, 39,
p. 10;i. Omitted are the numerous variations such as: Chameleo (Bonnaterre ;
1789); Cameleo (Bosc:lS03); Camaeleo (Dumeril :1806) ; Chameleon (Fleming:
1822); Chaelio (A. Smith :1831); Chamaeleon (Fitzinger :1S43) ; Camaehon
(DumSrll & Bibron :1863) ; some intentional emendations, others due to care
lessuess.

LOVERIDQE: E. AFRICAN REPTILES AND AMPHIBIANS 197

186.5. Pterosa^lrus Gray, Proc. Zool. Soc. London for 1864, pp. 467, 473.

Type by monotypy: Chamaeleo cristatus Stutchbury.
1865. Mierosaura^'^ Gray, Proc. Zool. Soc. London for 1864, pp. 467, 473.

Type by monotypy: M. melanocephala Gray.
1865. Phitvianola Gray, Proc. Zool. Soc. London for 18G4, pp. 467, 474.

Type by monotj-py: Chamaeleo namaquensis A. Smith.
1865. Lophosaiira Gray, Proc. Zool. Soc. Loudon for 1864, pp. 468, 474.

Type by restriction: Lacerta pumila Gmelin.
1865. Calumma Gray, Proc. Zool. Soc. London for 1864, pp. 468, 476.

Type by monotypy: Cliamaeleo cucullatus Gray.
1865. Crassonota Gray, Proc. Zool. Soc. London for 1864, pp. 468, 477.

Type by monotypy: Chamaeleo nasntus Dumeril & Bibron.
1865. Ensirostris Gray, Proc. Zool. Soc. London for 1864, pp. 468, 478.

Type by monotypy: E. melleri Gray.
1865. Sauroceras Gray, Proc. Zool. Soc. London for 1864, pp. 468, 478.

Type by monotypy: Chamaeleo rhinoceratus Gray.
1865. Dicranosanra Gray, Proc, Zool. Soc. London for 1864, pp. 468, 478.

Type by restriction : Cliamaeleo hif circus Gray = C. bifidtis

Brongniart.
1865. Cyneosaura Gray, Proc. Zool. Soc. London for 1864, pp. 468, 479.

Type by monotypy: Chamaeleo pardalis Cuvier.
1865. Calyptrosaura Gray, Proc. Zool. Soc. London for 1864, p. 468.

Type by present designation : Chamaeleo calyptratus A. Dumeril.
1865. Erizia Gray, Proc. Zool. Soc. London for 1864, p. 471. Type by

present designation: Chamaeleo senegalensis Daudin.
1865. Bilepis Gray, Proc. Zool, Soc, London for 1864, p. 472, Type by

tautonomy: Chamaeleo dilepis Leach.
1863. Archaius Gray, Proc. Zool. Soc. London for 1864, p. 475. Type by

present designation: Chamaeleo tigris Kuhl.
1956. Bicuspis Loveridge, Breviora (Cambridge, Mass.), No. 59, p. 2.

Type by original designation: Ehampholeon marshalli Bou-

lenger.^®

Chamaeleo senegalensis senegalensis Daudin
Senegal Chameleon

1802, Chamaeleo senegalensis Daudin, Hist. Nat. Eept., 4, p. 203 : Eegion
watered by the Senegal and Niger Rivers; Gambia and Guinea.

3& Microsanra. Under this name FitzSimons (1943: Transvaal Mus. Mem., 1,
p. 158), following Power (1932: Proc. Zool. Soc. London, p. 209) who. however,
used Lophosaura. would grant generic status to a group of South African
chameleons of which the best known is pumilua. If this course is followed,
then Bradypodion {Bradypodium of Gray) Fitzinger takes precedence over Micro-
saura, for Fitzinger definitely states : "Typus. Cham, pumilua Latr." Tentatively
I prefer to regard the group as subgeneric, but those who differ should removf
these three genera from the synonymy of Chamaeleo.

39 A prehensile-tailed species, but with the bicuspid claws of a Rhampholeon;
occupying an intermediate position between the two genera, hence Bicuspis..

198 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1863. Chamaeleo laevigatus Gray, Proc. Zool. Soc. London, p. 95: Five

hundred miles south of Khartoum, Sudan.

1864. Chamacleon scnegalensis var. Vciocephahis Gray, Proc. Zool. Soc.

London, p. 471 : Type in British Museum. No locality.

1887. Chamacleon sphaeropholis Reiclienow, Zool. Anz., 10, p. 370:

Kagera, west of Lake Victoria, Tanganyika Territory.

1951. Chamacleon scnegalensis v. Tihatiensis Monard, Mem. Inst. Franc.

Afrique Noire (Sci. Nat.), No. 1, p. 141: Tibati, French

Cameroon.
Range. Sudan, east to Eritrea, south through Ethiopia to
Uganda ; Kenya Colony and western Tanganyika Territory. Re-
corded from Angola and the Belgian Congo, northAvest to Sierra
Leone (M.C.Z. material) and Senegal.

Chamaeleo senegalensis anchietae Boeage*"
Angola Chameleon
1872. Chamneleo Anchietae Boeage, Jom. Sci. Lisboa, 4, p. 72, fig. — :

Huila, Mossamedes, Angola.
1950. Chamaeleo {Apola) vincTcci Laurent, Eevue Zool. Bot. Afr., 43,
p. 349: Swampy grassland sources of the Lofoi Eiver at 1750
metres, near Kundelungu Plateau Research Station, Katanga.
Belgian Congo.

1952. IChamaeleo anchietae mertensi Laurent, Revue Zool. Bot. Afr. 46,

p. 18: Muneshele, 1400 metres, Fizi Territory, Kivu, Belgian
Ruanda-Urundi.
1952. '^.Chamaeleo anehietae marmiguensis Laurent, Eevue Zool. Bot. Afr.,
46, p. 19 : Kipiri, 2000 metres, Marungu Plateau, Baudouinville
Territory, Tanganyika Province, Belgian Congo.
Range. Highlands of southern Tanganyika Territory, west
through highlands of eastern and southern Belgian Congo to
Angola.

Chamaeleo gracilis gracilis Hallowell Graceful Chameleon
1842. Chamaeleo gracilis Hallowell, Journ. Acad. Nat. Sci. Philadelphia,

8, p. 324, pi. xviii: Liberia.
1856. CJiamaeleo granulosus Hallowell, Proc. Acad. Nat. Sci. Philadel-
phia, p. 147 : West Africa.
1856. Chamaeleo Burchelli Hallowell, Proc. Acad. Nat. Sci. Philadelphia,
p. 147 : Fernando Po.

•10 1 have not seen mertensi and marungensia which were based on scanty
material (a ^ , $ and juvenile of the former, two $ $ of the latter), but I have
compared a $ paratype of vinckei (M.C.Z. 53262) with a Tanganyika 9
(M.C.Z. 31186) and failed to find differences that would justify its description.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 199

1885. Chamaeleo (Chaviacleo) Sirnoni Bocttger, Ber. Offenbach Ver.
Naturk., p. 175, footnote: Mountains of Ashanti, Gold Coast.

Range. French Somaliland, south through Uganda and Kenya
Colony to northern Tanganyika Territory (Longido and Mern
Mountains), west through tlie Belgian Congo (? and Angola)
where it meets with the race eUennei Schmidt (characterized by
spurless S S ) found around Banana; northwest (though not
definitely reported from the French Congo; Nigeria and Da-
liomey) to Senegal.

Chamaeleo dilepis roperi Boulenger*^
, Spurle.ss Flap-necked Chameleon
1890. Chamaeleon roperi Boulenger, Proc. Zool. Soc. London, p. 85,

pi. viii, fig. 4: Kilifi, north of Mombasa, Kenya Colony.
Range. Eastern Kenya Colony (Meru to Tana River) and
northeast Tanganyika Territory (around Kilimanjaro Mtn.
only ) .

Chamaeleo dilepis quilensis Bocage*- (doubtfully distinct)
18(3(5. Chamaeleo dilepis var. Quilensis Boeage, Jorn. Sci. Lisboa, 1, p-

59 : Rio Quilo, north of Cabinda, Cabinda.
1887. Chamaeleon parvilohus Boulenger, Cat. Lizards Brit. Mus., 3, p.

449, pi. xxxix, fig. 5: Natal; French Congo; Cameroon.
Range, (in part, being based solely on material in the M.C.Z.
from ) Kenya Colony ; Tanganyika Territory ; Southern Rho-
desia; Transvaal; Zululand; Natal; Southwest Africa; Angola:
Belgian Congo ; French Congo ; French Cameroon.

Chamaeleo dilepis dilepis Leach*"
Spurred F]ai)-necked Chameleon

11 Distinguished only by the ^ ^ being spurless like the $ J ; being identifi-
able only on the basis of adult ^ ^ , records require careful screening and the
range working out with considerable care.

■12 As this form, whose ^ ^ are usually spurred, is separable from d. dilepis
solely on the small size of its occipital flaps, subadult d. dilepis are apt to be
reported as d. quilensis with resulting confusion. M.C.Z. material reveals both
forms as present in five widely scattered countries. While a good series from a
given locality is likely to 1)« readily assignable to one form or the other, solitary
specimens are often intermediate in the degree of flap development. That we ai-"e
dealing with two sibling species seems improbable. FitzSimons (194.3, Transvaal
-Mus. jlem. No. 1, pp. 155-156) resolves that dilemma by according quilensis
varietal rank in order to indicate that it is something less than a subspecies. His
volume should be consulted for records of both forms south of the Zambezi.

43 Other apparently valid races occurring on the periphery of British East
Africa, and approached by individuals within the territories, are :
Chamaeleo dilepis petersii Gray, 1864. of Mozambique.
Chamaeleo dilepis isabellinus Giinther, 1893 (1892), from the Shire High-

lamls of Nyasaland.
Chamaeleo dilepis ruspolii Boettger, 1893, from Ogaden, Somalia.

200 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1819. Chamaeleo dilepis Leach, in Bowdich, Miss. Ashantee, App., p.

493 : French Congo.

1820. Chamaeleo bilobus Kuhl, Beitr. Zool. Anat., 1, p. 104: French

Congo.
1820. Chamaeleo planiceps Merrem, Vers. Syst. Amph., p. 162: Africa.
1864. Clmmaeleon petersii var. MrTcii Gray, Proc. Zool. See. London, p.

470 : East Africa.
1866. Chamaeleo Capellii Bocage, Jorn. Sci. Lisboa, 1, pp. 42, 59 : Ben-

guela, Angola.
1903. Chamaeleo anguslicoronatus Barbour, Proc. Biol. Soc. Washington,

16. p. 61 : Zanzibar Island.

Range, (in part, being based solely on material in the M.C.Z.
from) Kenya Colony; Tanganyika Territory; Pemba and Zanzi-
bar Islands ; Nyasaland ; Northern Rhodesia ; Southern Rhodesia ;
Cabinda ; Belgian Congo.

Chamaeleo bitaeniatus bitaeniatus Fischer"
Side-striped Chameleon
1884. Chamaeleo bitaeniatus Fischer, Jahrb. Hamburg, Wiss. Anst., 1.

p, 23, pi. ii, figs. 7a-b: Lake Naivasha, Kenya Colony.
1887. Chamaeleo bivittatus (lapsus: nomen nudum) F. Miiller, Yerh.

Naturf. Ges. Basel, 8, p. 294: Witu, Kenya Colony.
Range. Ethiopia and Somalia, south through Uganda and
Kenya Colony to northern and western Tanganyika Territory.

Chamaeleo bitaeniatus ellioti Gtinther
Montane Side-striped Chameleon
1895. Chamaeleon Ellioti Giinther, Ann. Mag. Nat. Hist., (6) 15, p. 524,

pi. xxi, fig. A : Bugoye, east foot of Euwenzori Mountains,

Uganda.
1912. Clmmaeleon bitaeniatus graueri Sternfeld, Sitzb. Ges. Naturf.

Freunde Berlin, p. 380, pi. xv, figs. 19-21, pi. xvii, fig. 33:

Bugoie and Eugege Forests above 2000 metres, Belgian Euanda-

Urundi.
1912. Chamaeleon bitaeniatus tornieri Sternfeld, Sitzb. Ges. Naturf.

Freunde Berlin, p. 383, pi. xvii, fig. 35 : Lendu Plateau, Ituri

District, Belgian Congo.

44 Owing to eesual or individual variation, members of the bitaeniatus group
are exceptioually difficult to place. Most large series are likely to contain one
or more individuals strikingly different from the rest, and such outstanding speci-
mens are apt to receive names. In addition to those listed, or tentatively
synonymized. here, two others have recently been described from neighbouring
territories. These are C. b. kinetensis K. P. Schmidt (1943) from Mount Kineti
at 10,458 feet, Imatong Mountains, southern Sudan, which appears to be inter-
mediate between typical bitaeniatus and b. ellioti; also C. b. schoutedeni Laurent
(19r>2) from Kabumbe Valley at 2400 metres, Kabobo Mountain, Albertville Ter-
ritory, Tanganyika Province, Belgian Congo.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 201

1922. Chamaeleon Bequaerti Witte, Revue Zool. Bot. Afr., 10, p. 69, pi.
ii, fig. 1 : Beni, Kivu District, Belgian Congo.

Range. Mountains (usually below 9000 feet) of western
Uganda ; Belgian Ruanda-Urundi and the adjacent Belgian
Congo.

Chctmaeleo bitaeniatus rudis Boulenger
Ruwenzori Side-striped Chameleon
1906. Chmnacleon rudis Boulenger, Ann. Mag. Nat. Hist., (7) 18, p. 473:

Ruwenzori Mountains above 10,000 feet, Uganda.
1983. Chamaeleon Burgeoni Witte, Revue Zool. Bot. Afr., 24, p. 120,
fig. — : Mombasa, near Lubero, Kivu District, Belgian Congo.
Range. Mountains (usually above 9000 feet) of western Ugan-
da ; Belgian Ruanda-Urundi and the adjacent Belgian Congo.

Chamaeleo bitaeniatus schubotzi Sternfeld
Kenya Side-striped Chameleon
1912. Chamaeleon hitaeniatus schuhotzi Sternfeld, Sitzb. Ges. Naturf.
Freunde Berlin, p. 381, pi. xv, fig. 24; pi. xvii, fig. 34: Mount
Kenya, 14,000 feet, Kenya Colony.
Range. Alpine zones of Mounts Kinangop and Kenya, Kenya
Colony ; Mounts Kilimanjaro and Nguru, Tanganyika Territory.

Chamaeleo bitaeniatus hohnelii Steindachner
Iligli-casqued Chameleon

1891. Chamaeleon Hohnelii Steindachner, Sitzb. Akad. Wiss. Wien., 100,
Abt. 1, p. 309, pi. i, figs. 1-la: Laikipia, 6000 feet, Kenya
Colony.

1891. Chamaeleon leikipiensis Steindachner, Sitzb. Akad. Wiss. Wien,
100, Abt. 1, p. 311, pi. i, figs. 2-2a: Laikipia, 6000 feet, Kenya
Colony.

1912. Chamaeleon bitaeniatus hergeri Sternfeld, Sitzb. Ges. Naturf.
Freunde Berlin, pp. 380-381, pi. xiv, fig. 16; pi. xvi, fig. 31:
"Sirgoi nordl. von Ravine," i.e. Sergoit, north of Eldama
Ravine Station, Kenya Colony (erroneously given as "Sirgoi
siidlich von Ravine," in a later paper of 1912).

Range. Highlands (usually below 9000 feet) of eastern
Uganda and Kenva Colony.

Chamaeleo bitaeniatus altaeelgonis Loveridge
Alpine High-casqued Chameleon
1935. Chamaeleon hitaeniatus altaeelgonis Loveridge, Bull. Mus. Comp.
Zool., 79, p. 15: Kaburomi, 10,500 feet, Mount Elgon, Uganda.

202 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Range. Alpine zoue (above 10,000 feet) of Mount Elgon,
Uganda and Kenya Colony.

Chamaeleo goetzei goetzei Tornier
Tanganyika Goetze Chameleon
1899. Ckamaeleon Goetzei Tornier, Zool. Anz., 22, p. 413, pi. ii, fig. 3:

Uhehe, Tanganj-ika Territory.
Range. Uzungwe ; Ubena ; Ukinga ; Rungwe and Poroto Moun-
tains, Tanganyika Territory.

Chamaeleo laterispinis Loveridge Spiny-sided Chameleon

1932. Chnmachon laterispinis Loveridge, Bull. Mus. Comp. Zool., 72,
p. 381 : Kigogo, 6000 feet, Uzungwe Mountains, Tanganyika
Territory.

Range. Uzungwe Mountains, Tanganyika Territory.

Chamaeleo incornutus Loveridge Ukinga Hornless Chameleon
1932. Chamaclcon incorniituH Loveridge, Bull. Mus. Conip. Zool., 72, p.
380: Madehani, 7000 feet, Ukinga Mountains, Tanganyika Terri-
tory.
Range. TTkinga ; Rungwe and Poroto Mountains, Tanganyika
Territory.

Chamaeleo tempeli Tornier Tubercle-nosed Chameleon

1899. Ckamaeleon Tempeli Tornier, Zool. Anz., 22, p. 411, pi. ii, figs.

2, 4: Uzungwe Mountains, Uhehe, Tanganyika Territory.

1900. Ckamaeleon tempeli var. ivolffi Tornier, Zool. Jahrb. Syst. 13, p.

613, fig. G (captions reversed and an 'f dropped from first
wolffi) : "Tardalla, " i.e. Tandala, Ukinga Mountains, Tan-
ganyika Territory.

Range. Uzungwe ; Ubena and Ukinga Mountains, Tanganyika
Territory.

Chamaeleo melleri (Gray) Giant One-horned Chameleon
1864. Ensirostris melleri Gray, Proc. Zool. Soc. London, p. 478, pi. xxxii,
fig. 1: "Mountains in the interior of East Africa" (as collected
by Dr. Meller of Zomba, restricted by Loveridge (1953) to
Zomba Mountain, Nyasaland).
Range. Savanna Forests of Tanganyika Territory and Nyasa-
land.

Chamaeleo spinosus Matschie Rosette-nosed Chameleon

1892. Ckamaeleon spinosus Matschie, Sitzb. Ges. Naturf. Freunde Berlin,
p. 105: Derema, Usambara Mountains, Tanganyika Territory.

LOVERIDOE: E. AFRICAN REPTILES AND AMPHIBIANS 203

Range. Mrgin Forests of Usambara Mountains, Tanganyika
Territory.

Chamaeleo tenuis Matschie Single Soft-horned Chameleon
1892. Chamaelcon tenuis Matschie, Sitzb. Ges. Naturf. Freiinde Berlin,

p. 106; Dererna, Usambara Mountains, Tanganyika Territory.
Range. Virgin Forests of Usambara Mountains, Tanganyika

Territory.

Chamaeleo xenorhinus Boulenger
Single Welded-horn Chameleon
1901. Chamaeleon xenorhinus Boulenger, Proc. Zool. Soe. London, 2. p.

135, pi. xii: Ruweuzori Mountains, 6000 feet, Uganda.
1929. Chamaeleo carpentcri Parker, Ann. Mag. Nat. Hist., (10) 3. p.

280, fig.: Ruwenzori Mountains, 6500 feet, Uganda.
Range. Virgin Forests (above 6000 feet) of Ruwenzori Moun-
tains. Uganda.

Chamaeleo fischeri excubitor Barbour

Kenya Hornless Chameleon
1911. Chamaeleo tenuis excubitor Barbour, October 31, Proc. Biol. Soc.

Washington, 24, p. 219: Meru District, Mount Kenya, Kenya

Colony.
1911. Chamaeleon affinis embucnsis Lonnberg, November 16, Svenska

Veteusk.-Akad. Handl., 47, No. 6, p. 19, pi. ii, fig. 3 : Forest on

eastern slopes of Mount Kenya, one day's march from Embu

Boma, Kenya Colony.
Range. Virgin forests of Mount Kenya, Kenya Colony.

Chamaeleo fischeri uthmolleri Miiller
Hanaug Hornless Chameleon'*^
1938. Chamaeleo uthmdlleri Miiller, Zool. Anz., 122, p. 20, figs. 1-2:
Mount Hanang (Gurui or Guruwe) at 2300 feet, central Tan-
ganyika Territory.
Range. Virgin forest of Mount Hanang, between Mkalama and
Kondoa Irangi, Tanganyika Territory.

Chamaeleo fischeri tcrvetanus Steindachner
Kilimanjaro Two-horned Chameleon
1891. Chamaeleon tavetanus Steindachner, Anz. Akad. Wiss. Wien, 28.

p. 142 : Taveta Forest, Kenya Colony.
1891. Chamaeleon tavetensis Steindachner, Sitzb. Akad. Wiss. Wien,
100. Abt. 1, p. 312, pi. i, figs. 3 3a: Taveta Forest, Kenya Colony.
^5 Assuming that the sexlng of the only known specimen is a ^ as stated.

204 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1891. Chamaeleo abbotti Stejneger, Proc. U. S. Nat. Mus., 14, p. 353,
fig. — : Mount Kilimanjaro, 4500 feet, Tanganyika Territory.

1893. Chamaeleo taitensis "Steind." (sic) Stejneger (lapsus for tave-
tanus), Proc. U. S. Nat. Mus., 16. p. 724.

Range. Virgin forests of Teita Mountains, Kenya Colony,
west to Kilimanjaro, Tanganyika Territory.

Chamaeleo fischeri multituberculatus Nieden
Western Usambara Two-horned Chameleon
1913. Cliamaeleon fischeri multituberculatus Nieden, Sitzb. Ges. Naturf.
Freunde Berlin, pp. 244, 248, figs. 27-29, pi. xv, fig. 7; pi. xvi,
fig. 11: < ' Phillipshof bei Willielmstal, " i.e. Magamba, near
Luslioto, western Usambara Mountains, Tanganyika Territory.
1913. Chamaelcon fischeri werncri Nieden (not of Tornier), Sitzb. Ges.
Naturf. Freude Berlin, pp. 243, 248, figs. 19-26, pi. xv, fig. 6;
pi. xvi, fig. 10: Ambangula and Mlalo, western Usambara Moun-
tains, Tanganyika Territory.
Range. Virgin forests of western Usambara Mountains, Tan-
ganyika Territory.

Chamaeleo fischeri fischeri Reicheuow

Eastern Usambara Two-horned Chameleon
1887. Chamaelcon Fischeri Eeichenow, Zool. Anz., 10, p. 371: Nguru

Mountains, Tanganyika Territory.
1895. CMnmeleon matschiei Werner, Verb. Zool. Bot. Ges. Wien, 45, p.

192 : Usambara Mountains, Tanganyika Territory.
1902. Cliamaeleon tornieri Werner, Zool. Jahrb. Syst., 15, p. 417, pi.

xxiv: "Mozambique" (presumably in error).
1913. Cliamaeleon fischeri vosseleri Nieden, Sitzb. Ges. Naturf. Freunde
Berlin, pp. 241, 247, figs. 11-18, pi. xv, fig. 5; pi. xvi, fig. 9:
Amaui; Bulwa; Nguelo in Usambara Mountains; Magrotto
Mountain ; Tanga, Tanganyika Territory.*^
Range. Virgin forests of eastern Usambara Mountains, south
to Nguru Mountain, Tanganyika Territory.

Chamaeleo fischeri ulugxiruensis Loveridge
Uluguru Two-horned Chameleon
1957. Chamaeleo fischeri uluguruensis Loveridge, Tanganyika Notes and
Eecords, No. 43, p. 3: Kingokwa, Uluguru Mountains, Tan-
ganyika Territory.

Range. Virgin forests of the Uluguru Mountains, Tanganyika
Territory.

*6 Restricted as the author also lists Ukami, which is in the Uluguru Moun-
tains ; and TJsaraiuo, i.e. Dar es Salaam region, in error.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 205

Chomaeleo fiillebomi Tornier Poroto Three-horned Chameleon

1900. Chamaelcon filUeborni Tornier, Zool. Jahrb. Syst., 13, p. (314, fig.

H: " Nonde-Nike, " i.e. Konde and Nyika (tribes); also
"Ngosi- Oder Peroto-Berges um 2200 m." i..e. Ngosi or Poroto
Mountains, Tanganyika Territory.
Range. Ngosi Volcano, Poroto Mountains, Ukonde, Tangan-
yika Territory.

Chcanaeleo werneri Tornier Uzungwe Three-horned Chameleon
1899. Chamncleon Werneri Tornier, Zool. Anz., 22. p. 258, fig. 1:
' ' Maschona-Gebiet, ' ' later corrected to Uzungwe Mountains,
Uhehe, Tanganyika Territory.
1932. Chamacleon loerneri dahagae Loveridge, Bull. Mus. Comp. Zool.,
72, p. 379: Dabaga, Uzungwe Mountains, Tanganyika Territory'.
Range. Uluguru and Uzungwe Mountains, Tanganyika Terri-
tory.

Chomaeleo johnstoni johnstoni Boulenger
Ruwenzori Three-horned Chameleon

1901. Chamaelcon johnstoni Boulenger, Proc. Zool. Soe. London, 2, p. 136,

pi. xiii: Ruwenzori Mountains, 6000 feet, Uganda.

1911. Chamaeleo'n graueri Steindachner, Anz. Akad. Wiss. Wien, 48. p.

177: Mountains northwest of Lake Tanganyika, Belgian Congo.

1912. Chamaelean johnstoni afflnis Stemfeld (not Riippell), Wiss.

Ergebn. Deut. Zentral-Afrika-Exped. 1907-1908, 4. p. 262, pi.
vii, fig. 3 : Virgin forest behind boundary mountains on north-
west shore of Lake Tanganyika, Belgian Congo.
Range. Virgin forests of Uganda; Belgian Ruanda-Urundi
and the adjacent Belgian Congo.

Chomaeleo jocksonii Boulenger

Kikuyu Three-horned Chameleon
1896. Chamaelcon JaclcsonU Boulenger, Ann. Mag. Nat. Hist., (6) 17,
p. 376: "Uganda" (error), later amended to Kikuyu, near
Nairobi, Kenya Colony.
1903. Chamaelcon jacTcsoni var. vauerescecae Tornier, Zool. Jahrb. Syst.,

19, p. 176: Nairobi, 6000 feet, Kikuyu, Kenya Colony.
Range. Highlands of Kenya Colony and Tanganyika Territory.

Chomaeleo deremensis Matschie

Usambara Three-horned Chameleon
1892. Chamaelcon deremensis Matschie, Sitzb. Ges. Naturf. Freunde Ber-
lin, p. 103; Derema, Usambara Mountains, Tanganyika Terri-
tory.

206 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Range. Usambara Mountains, Tanganyika Territory.

INTRODUCED

Chamaeleo oustaleti Mocquard^' Malagasy Giant Chameleon
1894. Chamaeleon Oustaleti Mocquard, C. R. Soc. Philom. Paris, No. 9,
p. 3; and 1895 (1894), Bull. Soc. Philom. Paris, (8), 7. p. 113:
Betsileo and Diego Suarez, Madagascar.
Range. Madagascar; but introduced into Ngong Forest, near
Nairobi. Kenya Colony.

Genus BROOKESIA Gray

48

1865. Broohesia Gray, Proc. Zool. Soc. London for 1864, pp. 468, 476.

Type by monotypy : Chamaeleo supcrciliaris Kuhl.
1874. Itliam'pholeon Giinther, Proc. Zool. Soc. London, p. 443, fig. — .

Type by monotypy: Chamaeleo spectrum Buchholz.
1942. Evoluticauda Angel, Mem. Acad. Malgache, No. 36, pp. 154, 178.

Type by subsequent selection: Broohesia nasvs Boulenger.*^

Brookesia kerstenii robecchi (Boulenger)
Somalia Pigmy-Chameleon
1892. Ehampholeon robecchii Boulenger, Ann. Mus. Civ. Stor. Nat.
Genova for 1891, (2) 12. p. 13, pi. i, fig. 3: "Wuorandi," i.e.
Warandi, ca. 5° 42' N., 47° 38' E., Somalia.
1897. Uliampholcon maadcra Meek, Field Mus. Nat. Hist., Zool. Series,

1. p. 183: Mandera, British Somaliland.
Range. British Somaliland, south through Somalia to north-
ern Kenya Colony.

Brookesia kerstenii kerstenii (Peters)
Kenya Pigmy-Chameleon

1' On April 11, 19G1, Miss V. Hardy-Mason brought to the Coryndon Memorial
Museum a iriaut chameleon which she had captured alive at Ngong in an area
undergoing deforestation.

Tlie reptile, a male, men.sures 18% inches from end of snout to tip of tail, the
latter forming 10 inches of the total. The snout does not terminate in a horn,
but from tlie crown of the head between the eyes there rises steeply a helmet-
like casque with strongly compressed median ridge. From behind this, along
the spinal ridge and, to a lesser extent, along the prehensile tail, is a continuou.s
serrated ridge formed of spinelike conical scales. A. I..

■18 Though Rhampholcon is inseparable from Brookesia on the basis of the
external characters cited l)y Giinther and Boulenger, it may well rank as a sub-
genus on the basis of the anatomical differences described by Parker (1942, Bull.
Mus. Comp. Zool., 91, pp. 80-81, hg. 8). In this sense all East African short-
tailed chameleons are probably referable to the subgenus Rhampholeon whose
range is Tropical Africa and Madagascar.

io As Angel designated no type for his genus Evohiticauda, I suggest the
iucliulod Hrnokcuiii nasiis be regarded as th(> typt'.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 207

1868. Chamaeleo Eerstenii Peters, Monatsb. Akad. Wiss. Berlin, p. 449:

"Wanga," i.e. Wange, south of Mombasa, Kenya Colony.
Range. Northern Kenya Colony, south to Tanganyika Terri-
tory.

Brookesia nchisiensis Loveridge Pitless Pigmy- Chameleon
1953. Brookesia nchisiensis Loveridge, Bull. Mus. Comp. Zool., 110, p.
190, pi. iii, fig. 1: Nchisi Forest, 5000 feet, Nehisi Mountain,
Nyasaland.
Range. Virgin forests of southwestern Tanganyika Territory,
south to Nchisi Mountain, Nyasaland.

Brookesia brevicaudata (Matschie)
Bearded Pigmy-Chameleon

1892. Chamaeleon (BrooJcesia) brevicaudatus Matschie (19.viii.92, fide

publishers), Sitzb. Ges. Naturf. Freunde Berlin, p. 107: Derema,
Usambara Mountains, Tanganyika Territory.

1893. Ehaiiipholeon Boettgeri Pfeffer, Mitt. Naturhist. Mus. Hamburg

for 1892, 10. p. 7G, pi. i, figs, (i 7 : Bagamoyo, Tanganyika Terri-
tory.

Range. Coastal Tanganyika Territory.

Brookesia brachyura ionidesi Loveridge
Beardless Pigmy-Chameleon
1951. Brookesin ionidesi Loveridge, Bull. Mus. Comp. Zool., 106. p. 179:

Kilwa, Southern Province, Tanganyika Territory.
Range. Southeastern Tanganyika Territory.

Brookesia temporalis (Matschie) Pitted Pigmy- Chameleon
1892. Cliamaeleon (Brookesia) temporalis Matschie, Sitzb. Ges. Naturf.
Freunde Berlin, p. 108: Derema, Usambara Mountains, Tan
ganyika Territory.
Range. Virgin forests of Usambara Mountains, Tanganyika
Territory.

Brookesia spectrum boulengeri (Steindachner)
Spectral Pigmy-Chameleon
1911. Ehampholeon boulengeri Steindachner, Anz. Akad. Wiss. Wien, 48,
p. 178: Mountains northwest of Lake Tanganyika, Belgian
Congo.
Range. \'irgin forests of western Uganda and eastern Belgian
Congo.

208 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Family SCINCIDAE

Genus MABUYA Fitzinger

1815. Mabuya Eafinesque, Anal. Nat. (Palermo), p. 16. Nomen nudum.

1826. Mabuya Fitzinger (part), Neue Class. Kept., pp. 23, 52. Type
by tautonomy: M. dominicensis Fitzinger = Lacertus mubouya
Lacepede which Fitzinger lists as a synonym.

1826. Spondylurus Fitzinger, Neue Class. Eept., p. 23 only. Type by
monotypy: Seincus sloanii Daudin.

1830. Euprepis Wagler (part), Nat. Syst. Amphib., pp. 132, 161. Type
by subsequent designation: Seincus agilis Eaddi = LacerUis
mabouya Lacepede.

1839. Herinia Gray, Ann. Nat. Hist., 2, p. 332. Type by monotypy: E.
capensis Gray.

1843. Eutropis Fitzinger, Syst. Kept., p. 22. Type by original designa-
tion: Euprepes sebae Dumeril & Bibron = Seincus carinatus
Schneider (part) + Seineus multifasciatus Kuhl + Euprepes
mneularius Blyth.

1843. Trachylepis Fitzinger, Syst. Eept., p. 22. Type by original
designation: Euprepes savignyi Dumeril & Bibron = Seincus
quinquetaeniatus Lichtenstein.

1843. Oxytropis Fitzinger, Syst. Eept., p. 22. Type by original designa-
tion: Euprepes m,erremii Dumeril & Bibron = Tiliqua capensis
Gray.

1845. Xystrolepis Tschudi, Fauna Peruana, Herp., p. 44. Type by
monotypy: Trachylepis punctata Tschudi (cf. Dunn, 1936, Proc.
Acad. Nat. Sci. Philadelphia, for 1935, 87. p. 557).

1845. Copeglossu-m Tschudi, Fauna Peruana, Herp., p. 45. Type by
monotypy : C. cinctum Tschudi i= Lacertus mabouya Lac6pede.

1845. Chioninia Gray, Cat. Lizards Brit. Mus., p. 116. Type by present
designation: Euprepes delalandii Dumeril & Bibron.

1848. Elabites Gistel, Naturg. Thierr., p. ix. Substitute name for
Euprepis Wagler, 1830.

1925. Mabuiopsis Angel, Eept. et Batr. in "Voyage de Ch. Alluaud et
E. Jeannel en Afrique Orientale (1911-1912). Eesultats scienti-
fiques. Vertebrata." (Paris), 2, p. 21. Type by monotypy
Mabuia Jeanneli Angel = M. irregularis Lonnberg.

Mabuya quinquetaeniata obsti Werner
Tanganyika Five-lined Skink^°

50 Only females and young are five-lined and blue-tailed, the males are strik-
ingly different. Ttie number of midbody scale-rows enables this inhabitant of
rocky outcrops to be broken up into several races, however the change is so
gradual that the limits have to be somewhat arbitrarily delined. North of Uganda
one encounters typical M. a. quinquetaeniata (Lichtenstein). South of the Zam-
bezi is M. q. margaritifer (Peters), described from Tete, Mozambique (cf. Lover-
idge, 1953, Bull. Mus. Comp. Zool., 110. pp. 196-200, 308).

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 20!:)

1913. Mabuia obsti Werner, Jahrb. Hamburg. Wiss. Anst. for 1912, 30,

p. 43: Kwa Mtoro, Usandawi, Tanganyika Territory.
1917. Mahuia quinquetaeniata hildebrandti Sternfeld (not Peters: 1874),
Wiss. Ergebn. Dent. Zweiter Zentral-Afrika-Exped. 1910-1911.
1. p. 438, pi. x.\iv, fig. 3: "Taita," i.e. Tcita, Kenya Colony.
Range. Uganda; Kenya Colony; Tanganyika Territory and
Cliapnani Island, Zanzibar (Tornier :1900), south through Nyasa-
iand to the Zambezi.

Mobuya maculilabris maculilabris (Gray)^^
Speckle-lipped Skink.

1845. Euprepis maculilabris Gray, Cat. Lizards Brit. Mus., p. 114: West
Africa.

1866. Euprepes anchietae Bocage, Jorn. Sei. Lisboa, 1, p. 62: "Zaire,"
i.e. Congo Eiver.

1879. Euprepes noiabilis Peters, Sitzb. Ges. Naturf. Freunde Berlin,
p. 36: Chinchoxo, Cabinda; and Pungo Ndongo, Angola.

1912. Ulabum maculilabris major Sternfeld, Wiss. Ergebn. Deut. Zentral-
Afrika-Exped. 1907-1908, 4. p. 232: Central Lake Region.

1912. Mabuia vmculilabris var. Tcwidjiviensis Sternfeld, Wiss. Ergebn.
Deut. Zentral-Afrika-Exped. 1907-1908, 4. p. 233: "Kwidjwi"
i.e. Idjwi Island, Lake Kivu, Belgian Congo.

1912. Mabuia macidilabris var. xcauensis Sternfeld, Wiss. Ergebn. Deut.
Zentral-Afrika-Exped. 1907-1908, 4. p. 233: Wau Island, Lake
Kivu, Belgian Congo.

1912. Mabuia, maculilabris var. schubotzi Sternfeld, Wiss. Ergebn. Deut.
Zentral-Afrika-Exped. 1907-1908, 4, p. 233, pi. vi, fig. 3: Kisenyi,
Lake Kivu, Belgian Euanda-Urundi; and Fort Bcni, Belgian
Congo.

1912. Mabuia maculilabris var. graueri Sternfeld, Wiss. Ergebn. Deut.
Zentral-Afrika-Exped. 1907-1908, 4. p. 234, pi. vi, fig. 4: Kisenyi,
Lake Kivu, Belgian Ruanda-Urundi ; also the Aruwimi-Ituri
region and between the Lualaba River and Lake Tanganyika,
Belgian Congo.

1912. Mabuin maculilabris var. rohrbecki Sternfeld, Wiss. Ergebn. Deut.
Zentral-Afrika-Exped. 1907-1908, A. p. 234: Nguru Mountains:
also "Langenburg," i.e. Manda, Lake Nyasa, Tanganyika
Territory.

51 This species is mucla in need of critical study on a continental scale. M. in.
major, rlesignated as a subspecies by its author to distinguish it from the color
varieties he named, appears to be a recognizable race, but when one attempts to
define its range diflticulties arise. Seemingly one must accept a great many form?,
based on average size and color (subject to fading), or reject them resolutely.
In the Usambara Mountains and adjacent coast one meets with a robust ami
handsomely coloured form to which, rightly or wrongly, I have been applying
the name "comorcntiis Peters."

210 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1912. Mahuia maculilahris var. bergeri Sternfeld, Wiss. Ergebn. Deut.

Zentral-Afrika-Exped. 1907-1908, 4, p. 235: Dufile, Sudan.
Range. Chiefly savanna areas of the southern Sudan and ??
Somalia {fide Calal)resi :1915), south through Uganda; Kenya
Colony and Tanganyika Territory (for off-shore islands see
races), west through Northern Rhodesia to Angola north and
west to French Guinea.

Mabuya maculilabris comorensis (Peters) ^'
Comoro Speckle-lipped Skink
1854. Eu'prepes comorensis Peters, Monatsber. Akad. Wiss. Berlin, p.

619: "Anjuan, " i.e. Johanna Island, Comoro Islands.
1882. Euprepes angasijanus Peters, Reise nach Mossambique, 3. p. 73 :
Angasiya, Grand Comoro Island.

1913. Mahuia comorensis var. infraUnpata Boettger, in Voeltzkow, Reise

in Ostafrika, 3, p. 329: Europa Island, Mozambique Channel.
Range. Chiefly virgin forest, or recently deforested, areas of
coastal Kenya Colony (Mombasa), south through Tanganyika
Territory (Usambara and Uluguru Mountains) also Zanzibar,
Mafia, Comoro and Europa Islands to Nyasaland (gallery forest
along Ruo River, Mlanje Mountain). Possibly Northern Rho-
desia (Meruwantipa).

Mabuya maculilabris albotaeniata Boettger '^

Pemba Speckle-lipped Skink
1913. MahuUi albotaeniata Boettger, in Voeltzkow, Reise in Ostafrika,

3. p. 350, pi. xxiv, figs. 1-2: Pemba Island.
Range. Pemba Island.

Mabuya maculilabris boulengeri Sternfeld ^^
Lindi Speckle-lipped Skink
1911. Mahuia hoidengeri Sternfeld, Sitzb. Ges. Naturf. Freunde Berlin,
p. 248: Makonde Plateau, Lindi Province, Tanganyika Terri-
tory.
Range. On tree-trunks in southeastern Tanganyika Territory
(Kilwa to Mikindani, inland to Songea), also Nyasaland (Mtim-
buka near Fort Johnston) .

52 See preceding footnote. Possibly the synonyms shouhl be recognized.

53 Examination of a cotype (M.C.Z. ;'>.3o2S) and a dozen other specimens fron;
seven localities on Pemba, reveal that this is a constantly recognizable color
form ; this has led me to reverse my synonyniizing of it in 1928.

5^ Its distribution suggests that boulengeri may be a full species ; the excep-
tionally long tail is correlated with a more strictly arboreal life.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 211

Mobuya brevicollis (Wiegmann)

1837. Euprepes brevicollis Wiegmann, Arch. Naturg., 3, Abt. 1. p. 133:

* ' Abyssinia, ' ' i.e. Ethiopia.
1837. Euprepes j)yi'>'Jiocephalus Wiegmann, Arch. Naturg., 3. Abt. 1,

p. 133 : Ascliik Island, Red Sea, Arabia.
185G. Tiliqita burtoni Blyth, Journ. Asiatic Soc. Bengal, 24. p. 306:

' ' Somali Country. ' '
1893. Mabnya chanleri Stejneger, Proc. U. S. Nat. Mus., 16. p. 721:

Tana River, Kenya Colony.
1895. Mabuya tcssellata Anderson, Proc. Zool. Soc. London, p. 636,

pi. xxxvi, fig. 2: near Aden (inferred from title), Arabia.
1905. Mabuin Eothschildi Mocquard, Bull. Mus. Hist. Nat. (Paris),

II. p. 286: Endessa, 8° 40' N., 40° E., Ethiopia.
1915. Mabuya pulchra Matschie, Sitzb. Ges. Naturf. Freunde Berlin,

p. 29: "Lahadsch, " i.e. Lahej, near Aden, Arabia.
Range. Sudan, east through Ethiopia and Eritrea to Asehik
Island and Arabia, south through British Somaliland and So-
malia to Uganda ; Kenya Colony and Tanganyika Territory.

Mabuya plonifrons ( Peters ) ^°

1878. Euprepes (Euprepis) planifrons Peters, Monatsb. Akad. Wiss.

Berlin, p. 203, pi. ii, fig. 2 : " Taita, ' ' i.e. Teita, Kenya Colony.

1878. Euprepes (Euprepis) taitanus Peters, Monatsb. Akad. Wiss.

Berlin, p. 203, pi. ii, fig. 3: "Taita," i.e. Teita, Kenya Colony.

1911. Mabuia diesneri Sternfeld, Sitzb. Ges. Naturf. Freunde Berlin,

p. 248: Kibwezi, Kenya Colony.
Range. Ethiopia, east to British Somaliland and Somalia,
south through Kenya Colony to Tanganyika Territory. ^^

Mabuya megalura (Peters) Grass-top Skink

1878. Euprepes (Mabuia) megalura Peters, Monatsb. Akad. Wiss. Berlin,

p. 204, pi. ii, fig. 4: "Taita," i.e. Teita, Kenya Colony.
1884. Eumeces massaianus Fischer, Jahrb. Hamburg, Wiss. Anst., 1. p.

18, pi. ii, fig. 5: Lake Naivasha Highlands, Kenya Colony.
Range. Ethiopia and Somalia, south through Uganda; Kenya
Colony and Tanganyika Territory to Belgian Ruanda-Urundi
and the adjacent Belgian Congo.

Mabuya bayonii keniensis Loveridge Eastern Bayon's Skink ^^

35 Specimens from northeast Tanganyika Territory are typical ; in the re-
mainder of the Territory southwest to Nyamkolo, Lake Tanganyika, Northern
Rhodesia, they are intermediate towards perrotetli (Dum^ril & Bibron) as I have
discussed elsewhere (1937, Tanganyika Notes & Records, No. 43, p. 4).

56 East African skinks formerly referred to bayonii (Bocage), differ from that
Angola species in possessing tricarlnate scales. Possibly the eastern form extends
to the Belgian Congo (Laurent :1952) and northeastern Angola.

212 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1956. Mabuya bayonii Iccniensis Loveridge, Breviora (Cambridge,
Mass.), No. 59, p. 2: Northern Uaso Nyiro, Sotik, Kenya Colony.
Range. Kenj^a Colony and Tanganyika Territory.^®

Mabuya varia varia (Peters)^' Savanna Variable Skiuk
1867. Euprepes (Euprepis) varius Peters, Monatsb. Akad. Wiss. Berlin,

p. 20: Tete, Mozambique.
1867. Euprepes Olivieri var. albo-punctatus Bocage, Jorn. Sci. Lisboa, 1,

p. 223 : Benguela and Catumbela, Angola.
1869. Euprepes damaranus Peters, Ofvers. Kongl. Vet. -Akad. Forh., p.

660: Damaraland, Southwest Africa.
1869. Euprepes {Mabuya) laevigatus Peters, Monatsb. Akad. Wiss.

Berlin, p. 434: Gerlachshoop, north of Middleburg, Transvaal.

1871. Euprepes {Exiprepis) Isselii Peters, Monatsb. Akad. Wiss. Berlin,

p. 567: "Keren," i.e. Cheren, Bogos, Eritrea.

1872. Euprepes angolensis Bocage, Jorn. Sci. Lisboa, 4, p. 78: Bibala,

Mossamedes; also Dondo, Angola.
1874. Euprepes {Euprepis) hildebrandtii Peters, Monatsb. Akad. Wiss.

Berlin, p. 372, pi. — , fig. 4: "Barawa, " i.e. Brava, Somalia.
Range. Sudan, east to British Somaliland, south through
Uganda; Kenya Colony and Tanganyika Territory to Natal,
west to Southwest Africa, north to Boma, Belgian Congo.

Mabuya varia braiini Tornier'^* Ukinga Montane Skink

1902. Mabuia brauni Tornier, Zool. Jahrb. Syst., 15, p. 585: Ukinga

Mountains, Tanganyika Territory,
Range. Tanganyika Territory (known onW from the holo-

type).

Mabuya irregularis Lonnberg Alpine-meadow Skinli

1922. Mabuia {striata ? var.) irregularis Lonnberg, Arkiv. Zool. 14, No.

12, p. 4: Soy, Kenya Colony.

1923. Mabuia Jeanneli Angel, Bull. Mus. Hist. Nat. (Paris), 29, p.

490: Mount Kinangop, Aberdare Mountains, Kenya Colony.
Range. Alpine meadows of Uganda (Mount Elgon) and
Kenya Colony (Mounts Kinangop and Kenya).

5' The Museum of Comparative Zoology has a cotype of isselii with 3 (instead
of 1) frontoparietals. Parker's comments (1942, Bull. Mus. Comp. Zool., 91,
p. 84) notwithstanding, I think the status of hildebrandtii is more probably that
of a synonym than a full species. Its distribution scarcely permits of its being a
recognizable race. All the available material should be restudied.

58 I am now inclined to think this skink may be a race of bocagii related to
M. b. mlanjenaia Loveridge of Nyasaland.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 213

Mobuya striata striata (Peters) ^^ Common Two-striped Skiiik

1844. Tropidolepis77ia striatum Peters, Ber. Akad. Wisa. Berlin, p. 36:
Mozambique.

1849. Euprepcs punctatissimus A. Smith, Illus. Zool. S. Africa, Rept., pi.
xxxi, fig. 1: northeastern districts of Cape Province, South
Africa.

1849. Euprepes sunderallii A. Smith, Illus. Zool. S. Africa, Kept., App.,
p. 11: Interior of South Africa.

1864. Euprepes grantii Gray, Proc. Zool. Soc. London, p. 62: "South-
Eastern Africa, ' ' i.e. Zambezi region.

1869. Euprepes vari.egatus Peters, Of vers. Kongl. Vet.-Akad. Forh., p.
660 : Damaraland, Southwest Africa.

1869. Euprepes wahlhergii Peters, Ofvers, Kongl. Vet. Akad. Forh., p.

661 : Damaraland, Southwest Africa.
1869. Euprepes (Euprepis) Griitzneri Peters, Monatsb. Akad. Wiss.

Berlin, p. 433 : Gerlaehshoop, north of Middleburg, Transvaal.
Range. Ethiopia and British Somaliland, sonth through
Uganda; Kenya Colony and Tanganyika (also Peniba ; Zanzibar
and Mafia Islands) to Natal, west to Southwest Africa, possibly
north to the Belgian Congo.

Genus RIOPA Gray""

1839. Eiopa Gray, Ann. Nat. Hist., 2. p. 332. Type by subsequent desig-
nation: Laeerta punctata Linnaeus.

1839. Chiamela Gray, Ann. Nat. Hist., 2. p. 332. Type by subsequent
designation : C. lineata Gray.

1839. Hagria Gray, Ann. Nat. Hist., 2, p. 333. Type by monotypy: E.
vosmaerii Gray.®-"-

1839. Campsodactylus Dumeril & Bibron, Erpet. Gen., 5, p. 761. Type
by monotypy : C. lamarrei Dumeril & Bibron = Hagria vosmaerii
Gray.

1843. Liosoma Fitzinger (not of Brandt: 1834), Syst. Rept., p. 22.

59 This synonymy requires restudying and the range correlated with those of
-If. 8. ellenbergeri Chabanaud of Zambezia, and the various forms described from
Angola and Southwest Africa.

60 Treated by Boulenger (1887, Cat. Lizards Brit. Mus., 3, p. 220) as a
"Section" of the genus Lygosoma. Subsequently, by common consent, raised to
full generic status, an action confirmed by Malcolm A. Smith (1937, Rec. Indian
Mus., 39, p. 227). It is in this sense it is used here. More recently Mittleman
(1952, Smithsonian Misc. Coll., 117, No. 17, pp. 8-10, 18-21), in a survey of
Lygosomine genera, has suggested further subdivision to a quite unacceptable
degree. He divides Riopa into six full genera. Of the species listed here he would
retain only mabuiiformis and tanae. To Mochltis he refers fernandi, sundevallii,
modesta and pembana, reviving Eumecla for the reception of anchietae.

61 Scincus vosmaerii Cocteau. which Gray cites as a synonym, appears to be
only a manuscript name.

214 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Type by original designation : Eumeces viicrolepis Dumeril &
Bibron.

1843. Sphenosoma Fitzinger, S3'st. Rept., p. 23. Type by original desig-
nation : Lacerta jmnciata Linnaeus.

1843. Evgongylus Fitzinger, Syst. Eept., p. 23. Type by original designa-
tion: Euvieces oiypellii Dumeril & Bibron =: Lacerta rufescens
Shaw (part).

1864. Mochlm Giinther, Proc. Zool. Soe. London, p. 308. Type by mono-
typy: M. punctnlatus Giinther = Eumeces sundevallii A. Smith.

1870. Eumecia Bocage, Jorn. Sci. Lisboa, 3, p. 67. Type by monotypy:
E. anchietae Bocage.

1880. Sci)acontias Giinther, Ann. Mag. Nat. Hist., (5) 6. p. 235. Type
by monotypy : S. modestus Giinther.

1892. Lcpidoihi/ris Cope, Journ. Morph., 7. p. 233. Tj^pe by subsequent
designation: Tiliqita fernmidi Burton.

1952. Squamicilia Mittleman, Smithsonian Mise. Coll., 117, No. 17, p. 9.
Type hj original designation: Eumeces isodactylus Giinther.

1952. Tachiigia Mittleman, Smithsonian Misc. Coll., 117. No. 17, p. 10.
New name for Liosoma Fitzinger : 1843 (preoccupied).

Riopa mabuiiformis Loveridge Mabuya-like Skink

193.'>. Riopa ntabuiifonnis Loveridge, Bull. Mus. Comp. Zool., 79. p. 12:

Ngatana, Tana River, Kenya Colony.
Range. Kenya Colony (known only from the type series).

Riopa tanae Loveridge Tana-Delta Skink

1935. Eiopa tanae Loveridge, Bull. Mus. Comp. Zool., 79, p. 11: Kan.

near mouth of the Tana River, Kenya Colony.
Range. Coastal Kenya Colony.

Riopa fernandi (Burton) Red-and-blaek Skink

1836. Tiliqua fernandi Burton, Proc. Zool. Soc. London, p. 62: Fernando

Po, Gulf of Guinea.
1845. Plestiodon Harlani Hallowell, Proc. Acad. Nat. Sci. Philadelphia,

p. 170: Liberia.
1854. Euprepis striata Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p.

98: Liberia.

1883. Eupr.epes (Tiliqua) clegans J. G. Fischer (not of Peters :1854),

Oester-Progr. Akad. Gymn. Hamburg, p. 3, pi. — , figs. 12-15:
Sierra Leone.

1884. Euprepcs leoninus Fischer (new name for elegans Fischer, preoc-

cupied), Abhand. Nat. Ver. Hamburg, 8, p. 7.

1885. Tiliqua nigripes F. Miiller, Verh. Naturf. Ges. Basel, 7. p. 704:

Gold Coast.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 215

Range. Uganda and Belgian Ruanda-Uruudi, west through
Belgian Congo to French Guinea.

Riopa sundevallii sundevallii ( Smith )*"'-
Suudevall 's Writhiug-Skink
1849. Eximices (Eiopa) siinderallii A. Smith, Illus. Zool. S. Africa, E«pt.,
App., p. 11: Country eastward of Cape Colony, i.e. Natal, South
Africa.
1854. Eumeces afer Peters, Monatsb. Akad. Wiss. Berlin, p. 619:
Mozambique Island; Boror; Inhambane and Mossimboa, Mozam-
bique.
1862. Eumeces reticulatvs Peters, Monatsb. Akad. Wiss. Berlin, p. 23:

New Barmen, Hereroland, Southwest Africa.
1864. Mochlus pimctulatus Giinther, Proc. Zool. Soc. London, p. 308:

Zambezi Expedition.
1868. Eumeces perdicicolor Cope, Proc. Acad. Nat. Sci. Philadelphia, p.

317: Zanzibar Island.
1870. Euprepes (Senira) Dumerili Steindaclmer, Sitzb. Akad. Wiss.

Wien, 62, Abt. 1, p. 341, pi. iii, fig. 5: Zanzibar Island.
1898. Lt/gosonia ferrandii Boulenger, Ann. Mus. Civ. Stor. Nat. Genova

(2), 18, p. 718, pi. ix, fig. 2: Lugh, Somalia.
Range. Sudan, east to French Somaliland (and Arabia),
south through Uganda ; Kenya Colony ; Tanganyika Territory ;
Zanzibar and Mafia Islands to Natal, northwest through the
Transvaal to Southwest Africa, north to French Equatorial
Africa where one encounters intermediates with a western race."'^

Riopa modesta modesta (Giinther) Brown Writhing-Skink
1880. Sepacontias modestus Giinther, Ann. Mag. Nat. Hist., (5) 6.

p. 235 : Mpwapwa, Ugogo, Tanganyika Territory.
Range. Ethiopia (1895) and Uganda (1902), east and south

through Kenya Colony and Tanganyika Territory direct to

Nyasaland; also recorded from Belgian Ruanda-Urundi (1912).

the Belgian Congo (1952) and Angola (19'S7).'^

82 Omissions from the synonymy refer to the western subspecies.

63 /J. s. guinrensis (Peters :1879), erroneously recorded from East Africa hy
Tornier (1S96), is a barely recognizable form on the basis of fewer subdlgital
lamellae beneath the fourth toe. It (including chaperi Chabanaud :1SS4_) ranges
from French Efiuatorial Africa (intermediates) west to Portuguese Guinea. In
this connection the status of houyi (Sternfeld :1916) and mocquardi (Chabanaud :
1917) require investigating.

fi-i The resemblance to sundevallii of occasional examples of modesta make it
uecessar.v to view with suspicion the five solitary records of its occurrence outside
its custoniarv ran^re. British Somaliland records are referable to R. m. somalica
Parker (1942).

216 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Riopa pembana (Boettger) Pemba Island Writhing-Skink
1913. Lygosoma (Eiopa) pembanum Boettger, in Voeltzkow, Eeise in

Ostaf rika, 3, p. 350, pi. xxiv, figs. 4-5 : Pemba Island.
Range. Coastal Kenya Colony (probably introduced) and

Pemba Island.

Riopa anchietae (Bocage) Western Serpentiform Skink

1870. Eumecia anchietae Boeage, Joru. Sci. Lisboa, 3, p. 67, pi. 1:

Huilla Plateau, Mossamedes, Angola.
Range. Western Kenya Colony and Tanganyika Territory,

southwest to Northern Rhodesia and Angola, north to Belgian

Congo.

Genus LYGOSOMA Hardwicke & Gray""
1827. Lygosoma Hardwicke & Gray, Zool. Journ., 3, p. 228. Type by
monotypy: Lacerta serpens Bloch = Anguis quadrupes Linnaeus.

Subgenus LEPTOSIAPHOS Schmidt

1943. Leptosiaphos K. P. Schmidt, Field Mus. Nat. Hist. Zool. Ser., 24.
p. 332. Type by original designation: Lygosoma meleagris
Boulenger.

Lygosoma kilimense Stejneger^ Kilimanjaro Five-toed Skink
1891. Lygosoma Icilimensis Stejneger, Proc. U. S. Nat. Mus., 14, p. 405:

Mount Kilimanjaro, Tanganyika Territory.
1900. Lygosoma dathrotis Boulenger, Ann. Mag. Nat. Hist., (7) 6,

p. 194: foot of Mount Kenya, Kenya Colony.
1903. Lygosoma thomasi Tornier, Zool. Jahrb. Syst., 19, p. 175: Nairobi,

Kenya Colony.
1907. Lygosoma aloysii-sahaudiae Peracca, Boll. Mus. Zool. Anat. Comp.

Univ. Torino, 22, No. 553, p. 1: Mitiana and Toro, Uganda.

65 Departing from the custom employed elsewhere, I have omitted the lengthy
and quostionable synonymy of Lygosoma. Its East African representatives all
belong to the subgenus Leptosiaphos, separated by Schmidt from 8aiph08 (correct
spelling) of Gray :1831 whose range is restricted to southeast Asia and Australia.

Mittleman (1952, Smithsonian Misc. Coll., 117, No. 17, pp. 6, 20) regards
Leptosiaphos as a full genus, separating it from Saiphos on characters I have
had tested on 4 African species (177 specimens) and 6 Australian (22 speci-
mens), and I found the following to hold good.

Ear-opening present though often minute ; anal not or but slightly enlarged ;

range : East and Central Africa Leptosiaphos

Ear-opening absent ; anal noticeably enlarged ; range : Malaya ; Indonesia ;
Australia Saiphos

66 De Witte, who subsequently (1953) secured 182 of these skinks, concurs with
my action in synonymizlng dewittei. L. gromieri, referred by its author to Riopa,
had a longitudinally divided frontonasal and so simulated a pair of supranasals ;
its synonymizlng with kilimensis can be regarded as provisional.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 217

1925. 'iLygosoma gromieri Angel, Bull. Mus. Hist. Nat. (Paris), 31. p.
419: (On termitarium near station) Tsavo, Kenya Colony.

1933. Lygcysomu (Siaphos) coviprcssicauda Witte (not Werner: 1897),

Revue Zool. Bot. Afr., 23, p. 175, figs. 1-4: Sandoa, Belgian
Congo.

1934. Siaphos dewittei Loveridge (new name for compressicauda Witte,

preoccupied), Copeia, p. 184.
1943. Lygosoma (Lcptosiaphos) weberi K. P. Schmidt, Field Mus. Nat.
Hist. Zool. Scr., 24. p. 335, fig. 27: Lotti Forest at about 3300
feet on west side of Imatong Mountains, Sudan.
Range. Virgin forest, usually on mountains, from southern
Sudan through Uganda; Kenya Colony and Tanganyika Terri-
tory, west through the Belgian Congo to Congulu, Angola.

Lygosoma kutuense Lonnberg Kenya Five-toed Skink

r.ni. Lygosoma ]:utuent>is Lonnberg, Kungl. Svenska. Vetensk.-Akad.

HandL, 47, No. 6, p. 17 fig. 3: Near Kutu Village, between

Embu and Fort Hall, Kenya Colony.
Range. Kenya Colony (known only from the holotype).

Lygosoma graueri graueri Sternfeld Ruanda Five-toed Skink
19lL\ Lygosoma graueri Sternfeld, Wiss. Ergebn. Deut. Zentral-Afrika-
Exped. 1907-1908, 4, p. 240, fig. 3: Locality given only for races,
so suggest restricting it to bamboo forest at foot of Mount
Karisimbi, Belgian Ruanda-Urundi.
hH'2. Lygosoma graueri quvnquedigiiata Sternfeld, Wiss. Ergebn. Deut.
Zentral-Afrika-Exped. 1907-1908, 4. p. 241, fig. 3b, pi. vi, fig. 5:
Bamboo forest at foot of Mount Karisimbi, Belgian Euanda-
Urimdi.

Range. Virgin forests of Uganda ; Belgian Ruanda-Urundi
and the adjacent Belgian Congo.

Lygosoma meleagris Boulenger"' Ruwenzori Four-toed Skink

1907. Lygosoma meleagris Boulenger, Ann. Mag. Nat. Hist., (7) 19, p.

488: Mubuku Valley, 7000 feet, Ruwenzori Mountains, Uganda.

1932. Siaphos meleagris helleri Loveridge, Proc. Biol. Soc. Washington,

45, p. 113: Bugongo Ridge, 9500 feet, Ruwenzori Mountains,
Belgian Congo (not Uganda side).

1933. Lygosoma (Siaphos) Burgeoni de Witte, R^vue Zool. Bot. Afr.,

24, p. 116, figs. 1-2: Kalonge, 6725 feet, Ruwenzori Mountains,
Belgian Congo.
1941. Lygosoma meleagris haclcarsi de Witte, Inst. Pares Nat. Congo
Beige, Expl. Pare Nat. Albert, Miss. G. F. de Witte (1933-

67 Dr. de Witte (1953) concurs with the synonymy as given here.

218 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1935), 33. p. 138, figs. 39-40, pi. ii, figs. 4-6: Kamatembe, 6825
feet, Pare National Albert, Belgian Congo.
Range. Virgin forests of Uganda ; Belgian Ruauda-Urundi
and the adjacent Belgian Congo.

Genus ABLEPHARUS Fitzinger

6S

1823. Ahlepharus Fitzinger, in Lichtenstcin, Verz. Doul)l. Zool. Mus.
Berlin, p. 103. Type by monotypy: A. pannonicus Fitzinger.

1833. Lerista Bell, Proc. Zool. Soc. London, p. 99. Type by monotypy:

L. lineaia Bell.

1834. Cryptohlepharnm (sic) Wiegmann, Herp. Mexicana, p. 12. Type

by subsequent designation: Ahlepharris poecilopleuro (sic)

Wiegmann.
1839. Petia Gray, Ann. Nat. Hist., 2, p. 335. Substitute name for

" Cryptohlcpharus Wiegmann."
1843. Microblepharis Fitzinger, Syst. Kept., p. 23. Type by original

designation: Ahlepliarus menestriesii Dumeril & Bibron =

Scincus hivii tains Menestries.
1843. Ophiopsis Fitzinger, Syst. Eept., p. 23. Tj-pe by original designa-
tion: Lerista lincata Bell.
1845. Morethia Gray, Cat. Lizards Brit. Mus., p. 65. Type by monotypy:

M. anomahis Gray.
1845. Menetia Gray, Cat. Lizards Brit. Mus., pp. 65, 66. Type by

monotypy : M. greyii Gray.
1845. Miculia Gray, Cat. Lizards Brit. Mus., p. 66. Type by monotypy:

M. elegams Gray.
1872. Blepharosteres Stoliczka, Proc. Asiatic Soc. Bengal, p. 74. Type

by monotypy : B. grayanus Stoliczka.
1881. Phaneropis Fischer, Arch. Naturg., 47. Abt. 1, p. 236. Type by

monotypy: P. muelleri Fischer.

Ablephorus boutonii airicanus Sternfeld
Coral-rag Snake-eyed Skink
1918. Ablepharus boutoni africanus Sternfeld, Abhand. Senckenberg.
Nat. Ges., 36, p. 423 : Manda Island and Malindi, Kenya Colony ;
Pemba Island.
Range. Rocky coastal stretches of Somalia; Kenya Colony
(including Manda, Lamu and Mombasa Islands) ; and Tan-

68 Recently Mittlenian (19.52, Smithsonian Misc. CoU., 117. No. 17, pp. 13. 19),
following Stejneger, bas revived Crytoblepharua as a full genus on the basis of
its frontoparietal being fused with the interparietal, together with the size of
its ear-upHiiing. To uie this seems a wholly unnatural division (of. footnote to
A. wahlbergii). It is customary to cite "Herp. Mesicana" for Cryptohlephwrus,
but Wiegmann gives Cryptoblepharum as a synonym of Ablepharus without inten-
tion of proposing a new genus.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 219

ganyika Territory; also Pemba, Zanzibar and Mafia Islands.

Ablepharus wahlbergii (Smith)"" Savanna Snake-eyed Skink
1849. Cnjptohlcpharus icalilhergii A. Smith, Illus. Zool. S. Africa, R*pt.,

App., p. 10: Natal, South Africa.
1894. Ablephurus carsonii Bouleuger, Proc. Zool. Soc. Loudon, p. 735,
pi. xlLx, figs. 4-4a: "Fwambo," i.e. Fwamba, Northern Rhodesia.
1924. Ablepharus massaiensis Augel, Bull. Mus. Hist. Nat. Paris, 30.

p. 52: Masai Plains near Nairobi, Kenya Colony.
Range. Ethiopia and British Sornaliland, south through So-
malia, Kenya Colony and Tanganyika Territory (reported from
Grave Island, Zanzibar, but possibly in error for boutonii afri-
canus; also Mafia Island) to Natal, west to Southwest Africa,
northeast through Angola to Belgian Congo. Unknown from
I^ganda.

Ablepharus megalurus Nieden Blue-tailed Snake-eyed Skink
1913. Ablepharus megalurus Nieden, Mitt. Zool. Mus. Berlin, 7, p. 89:
"Kinjanganja in Turu, 4° 50' S." i.e. Unyanganyi, Turn,
Tanganyika Territory.
Range. Central Tanganyika Territory (from Unyanganyi
south to Saranda on the Central Railway).

Genus SCELOTES Fitzinger'°

1826. Scelotes Fitzinger, Neue Class, Rept., pp. 23, 53. Type by niono-
iypj: Bipes anguineus Merrem == Anguis bipes Linnaeus.

1830. Zygnls Wagler (not Oken:1816), Nat. Syst. Amphib., p. 160.
Type by monotypj': Anguis bipes Linnaeus.

1839. merinia Gray, Ann. Mag. Nat. Hist., 2. p. 332. Type by monotypy:

69 A. carsonii Is only tentatively referred to the synonymy. It was based on a
single skink whose interparietal was fused with the frontoparietal, probably an
individual aberration as wahlhergii is common at Abercorn, from which Pwamba
is not very distant. The status of A. tancrcdi Boulenger (1909) from Dabarif,
Ethiopia, and A. u-ilsani Werner (1914) from Talodi, Sudan, also merit investi-
gation as both are still known only from their respective holotypes.

70 Overlooking Gongylomorphus of Fitzinger, Witte & Laurent (1943, Mem.
Mus. Royal Hist. Nat. Belgique, (2) Ease. 26, p. 3) revived its synonym Thyrus
as a monotypic genus.

They described Proscelotes for the reception of two species (eggeli and ulu-

be regarded as a recognizable subgenus.

So far as this check list is concerned, Witte & Laurent would refer the two
forms of tetraduvtijlun to Scpsina which, together with Herpetosaura they raise
to full generic rank, while synonymizing Jlelanosepa with Scelotes. A multiplicity
of small genera emphasizes diflferences at the expense of relationships, though in
this instance the authors claim that similarities are the result of convergences.

220 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

H. capensis Gray, which therefore antedates Gongylus capensis

sp. n. of A. Smith: 1849 Ji
1839. AmpUglossus Dumeril & Bibron, Erpet. Gen., 5, pp. 606, 608. Type

by monotypy: A. astrolahi Dumeril & Bibron.
1843. Gongylomorphus Fitzinger, Syst. Bept., p. 22. Type by monotypy:

Scincus hojerii Desjardin.
1845. Thyrtis Gray, Cat. Lizards Brit. Mus., p. 124. Type by monotypy:

Scincus iojerii Desjardin.
1849. Lithophilus A. Smith (not of Fi-61ich:1799), Illus. Zool. S. Africa,

Rept., App., p. 12. Type by subsequent designation: L. inornatiis

A. Smith.
1854. Hfirpetosaura Peters, Monatsb. Akad. Wiss. Berlin, p. 619. Type

by monotypy: B.. arenicola Peters.
1861. Scpomorphus Peters, Monatsb. Akad. Wiss. Berlin, p. 422, Type

by monotypy: S. caffer Peters.
1866. Sepsina Boeage, Jorn. Sci. Lisboa, 1, p. 62. Type by monotypy:

S. angolensis Boeage.
1866. Dumerilia Boeage, Jorn. Sci. Lisboa, 1, p. 63. Type by monotypy:

D. bayonii Boeage.

1874. llhinoscincus Peters, Monatsb. Akad. Wiss. Berlin, p. 374. Type

by monotypy: Sepsina (Bhinoscincus) tetradactyla Peters.

1875. Scincodipus Peters, Monatsb. Akad. Wiss. Berlin, p. 551. Type

by monotypy: S. congicus Peters = Dumerilia bayonii Boeage.
1887. Herpetoseps Boulenger, Cat. Lizards Brit. Mus., 3. p. 416. Tj^je by

monotypy: E. anguineris Boulenger.
1943. Proscelotes Witte & Laurent, Mem. Mus. Eoyal Hist. Nat. Bel-

gique, (2) Fasc. 26, pp. 9, 13. Type by original designation:

Scelotes eggeli Toruier.

Scelotes eggeli Toruier Usambara Five-toed Scelotes

1902. Scelotes eggeli Tornier, Zool. Anz., 25, p. 700: Kwai, western

Usambara Mountains, Tanganyika Territory.
Range. Usambara Mountains Tanganyika Territory.

Scelotes uluguruensis Barbour & Loveridge
Uluguru Five-toed Scelotes
1928. Scelotes uluguruensis Barbour & Loveridge, Mem. Mus. Comp.

vi Fi-om the brief description. Dr. V. FitzSimous considers thiat H. capensis
may well be a Scelotes. Mr. J. C. Battersby writes me that the type of H.
capensis Gray, 1838, is uot to be found in the British Museum, and apparently
was not there in 1887 when Boulenger wrote the Cat. Lizards Brit. Mus., 3,
p. 209 and listed -"fHerinia" in the synonymy of Lygosoma. In 1935 Malcolm
Smith (Fauna Brit. India. Rept. Amph., 2, p. 216) transferred it to the synonymy
of Muhuya, possibly on the assumption that it was identical with Tiliqua
capensis Gray, 1830, which was also described from the Cape of Good Hope.
The latter species, as Mahuya capensis (Gray), 1830, is in general use for
Scincus trivittatus Cuvier, 1829, preoccupied by trivittatus Hardwicke & Gray.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 221

Zool., 50. p. 167, pi. iv, fig. 6: Bagilo, 5000 feet, Uluguru Moun-
tains, Tanganyika Territory.
Range. Uluguru Mountains, Tanganyika Territory.

Scelotes tetradactylus tetradactylus (Peters)
Eastern Four-toed Scelotes
1874. Sepsina (Ehinoscincus) tetradactyla Peters, Monatsb. Akad. Wiss.

Berlin, p. 374: "Zanzibar Coast," i.e. Tanganyika Territory.
Range. Eastern Tanganyika Territorj^ south to Nyasaland.

Scelotes tetradactylus hemptinnei (Witte)
Western Four-toed Scelotes
1933. Sepsitia Hemptinnei de Witte, Eevue Zool. Bot. Afr., 23. p. 188:

Lukafu, Kundelungu, Katanga, Belgian Congo.
Range. Near Ujiji, Tanganyika Territory, west to Belgian
Congo.

Genus MELANOSEPS Boulenger"

1887. Melanoscps Boulenger, Cat. Lizards Brit. Mus., 3, p. 422. Type
by monotj'py : Herpetosaura atra Giinther.

Melanoseps ater longicouda Tornier
Long-tailed Limbless-Skink
1900. Melanoseps ater var. longicaiula Tornier, Zool. Jahrb. Syst., 13,
p. 602: Masailand, and " Karagwe am Pangani, " i.e. Korogwe,
Pangaui River, Tanganyika Territory.
Range. Southern Kenya Colony and northern Tanganyika
Territory (whether the skink from Uluguru Mountains should
be included is uncertain).

Melanoseps ater rondoensis Loveridge
Rondo-Plateau Limbless-Skink
1942. Melanoseps ater rondoensis Loveridge, Bull. Mus. Comp. Zool.,
91. p. 360: Nchingidi, 2700 feet, Rondo Plateau, Lrndi District,
Tanganyika Territory.
Range. Rondo Plateau, Tanganyika Territory.

"2 To the synopsis of the races furnished by me (1942, Bull. Mus. Comp. Zool.,
91, p. 359) should be added the Nyasaland forms described (1943, Bull. Mus.
Comp. Zool., 110. pp. 220-223).

Witte & Laurent (1943, Mem. Mus. Royal Hist. Nat. Belgique (2), Fasc.
26, p. 32) may well he right in merging Melanoseps with Scelotes but the decision
merits careful revisionary study of the entire complex. These limbless skinks are
associated with primary forest, either montane or gallery.

222 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Melanoseps crter matengoensis Loveridge
Mateugo-Highlands Limbless-Skink
1942. Melanoseps atcr matengoensis Loveridge, Bull. Mus. Conip. Zool.,
91, p. 361 : Ugano, Matengo Highlands west of Songea, Tan-
ganyika Territory.

Range. Matengo Highlands, Tanganyika Territory.

Melanoseps ater uzxxngwensis Loveridge
Uzungwe-Mountains Limbless-Skink

1942. Melanoseps ater usungtcen.sis Loveridge, Bull. Mus. Comp. Zool.,

91, p. 361: Kigogo, Uzungwe Mountains, Tanganyika Territory.
Range. Uzungwe Mountains, Tanganyika Territory.

Genus SCOLECOSEPS Loveridge

1920. Scolecoseps Loveridge, Proc. Zool. Soc. Loudon, p. l.')9. Type
S. boulengeri Loveridge.

Scolecoseps acontias (Werner) Sandy-soil Limbless-Skink
1913. .]felanoseps aeontias Werner, 1912, Jalirb. Hamburg. Wiss. Anst.

for 1912, 30, p. 19: Dar es Salaam, Tanganyika Territory.
Range. Coastal (Dar es Salaam to Kilwa) Tanganyika Terri-
tory.

Genus ACONTIAS Cuvier

1817. Acontius Cuvier, Regne Animal, 2, p. 60. Type by monotypy:
Angtiis mcleagri^ Linnaeus.

Acontias percivali Loveridge Teita Mountains Limbless-Skink
19.15. Acontias percivali Loveridge, Bull. Mus. Comp. Zool., 79. p. 13:

Foot of Mount Mbololo, Teita Mountains, Kenya Colony.
Range. \'ieinity of Voi, Kenya Colony.

Genus FEYLINIA Gray

1845. Feylinia Gray, Cat. Lizards Brit. Mus., p. 129. Type by monotj-py:

F. currori Gray.
18.")6. Anehjirops A. Dumeril, Revue Mag. Zool., (2) 8, p. 420. Type by

monotypy: Acontias elega/ns Hallowell.
1857. Sphcnorhina Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p. 52.

Typo by monotypy: Acontias elegans Hallowell.

1943. Chahanamlia Witte & Laurent, Mem. Mus. Royal Hist. Nat.

Belgique, (2) Fasc. 26, p. 37. Type by original designation:
Feylinia boiiU tifferi Cliabanaud.

'n

LOVEBIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 223

Feylinia currori elegotis (Ilallowell)'^
Western-Forest Limbless-Skink
1852. Acontias clegans Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p.

65 : " Liberia, ' ' probably in error for Gabon.
Range. U<ian(la (Entebbe; Sesse Islands, etc.) and western
Tanganyika Territory (Bnkoba). west throufrh the Belgian and
French Congo.

Family CORDYLIDAE (inc. GERRHOSAURIDAE) '^
Genus CORDYLUS Laurenti

1763. Cordylus Gronovius, Zoophyl. Gronov. . . ., p. 13. (A work
specifically rejected for taxonomic purposes in rule 89 of the
Internat. Comm. Zool. Nomencl. • — cf. 1926, Proc. Biol. Soc.
Washington, 39. p. 103).

1768. Conlyhis Laurenti (part), Syn. Eept., p. 51. Type by devious
tautonomy: C. verus Laurenti := Lacerta cordylus Linnaeus.

1820. Zonurus Merrem, Vers. Syst. Amphil)., p. 57. Type by monotypy:
Lacerta cordylus Linnaeus.

Cordylus cordylus tropidostemum (Cope)
Eastern Spiny-tailed Lizard
1869. Zonurus iropidosternum Cope, Proc. Amer. Philos. Soc, 11, p. 169:

"Madagascar" (presumablj' error for Mozambique).
1889. Zonurus frenatus Pfeffer, Jahrb. Hamburg. Wiss. Anst., 6, p. 6:

Mhonda, Eastern Province, Tanganyika Territory.
1934. Zonurus parlccri Cott, Proc. Zool. Soc. London, p. 151, pi. ii:
Auiatongas, Mozambique.

Range. Coastal Kenya Colony (near Malindi), south through
Tanganyika Territory ; Mozambique and Nyasaland to Southern
Rhodesia (where it meets with C. c. jonesii and C. c. rhodesi-
anus) .

Cordylus uldngensis (Loveridge) Ukinga Spiny-tailed Lizard
li)32. Zonurus u}:in(;ensis Loveridge, Bull. Mus. Comp. Zool., 73. p. 378,

pi. iii, fig. 2: Tandala, Ukinga, Tanganyika Territory.
Range. Southern Highlands of Tanganyika Territory.

73 Witte and Laurent (1943) make this a full species. The genus is in great
need of couipreheiislve revisifm based on all available material. Our Entebbe
specimen has the ocular in contact with the second labial ; also 26 midbody
scale-rows.

74 McDowell and Bogert (19.54, Bull. Am. Mus. Nat. Hist., 105, pp. 98-102) have
advanced reasons for the merging of these two families. For synonymy of family
names see Loveridge, 19.57, Tanganyilia Notes *: Records, No. 4.S, p. ti.

224 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Genus GERRHOSAURUS Wiegmann

1828. Gerrhosaurus Wiegmann, Isis von Oken, p. 378. Type by mono-
typy: G. flavigularis Wiegmann.

1836. Pleurotuchus A. Smith, Mag. Zool. Bot. (Jardine), 1. p. 142. Type

by monotypy: P. typicns Smith.

1837. Pleurotychus Wiegmann, Arch. Naturg., 3, Abt. 2, p. 228. Emenda-

tion for Pleurotuchus A. Smith.

Gerrhosaurus major bottegoi Prato"
Eritreau Great Plated-Lizard
1895. Gerrhosaurus bottegoi del Prato, Atti, Soc. Ital. Sci. Nat.

(Milano), 35, p. 19, figs. 1-la: Ghinda Valley, Eritrea.
Range. Eritrea, south through Ethiopia and Somalia to Kenya
Colony and central Tanganyika Territory.

Gerrhosaurus major major Dumeril
Zanzibar Great Plated-Lizard
1851. Gerrhosaurus Major A. Dumeril, Cat. Method. Coll. Rept. Mus.

Paris, p. 139: Zanzibar Island.
1889. Gerrhosaurus zanzibaricus Pfeffer, Jahrb. Hamburg. Wiss. Anst.,

6, p. 7: "Kibueni," i.e. Kibweni, Zanzibar Island.
1906. Gerrhosaurus bergi Werner, Zool. Anz., 30. p. 54, figs. 1-3: Usam-

bara Mountains, Tanganyika Territory.
Range. Eastern Kenya Colony; Tanganyika Territory and
Zanzibar Island, south to northern Mozambique (at Lumbo and
Tete).

Gerrhosaurus major grandis Boulenger^®
Zululaud Great Plated-Lizard
1908. Gerrhosaurus grandis Boulenger, Ann. Natal Mus., 1, p. 225, pi.

xxxvi: Zululand, Natal.
Range. Central (Morogoro) and southeast (Kilwa; Liwale;
Tunduru) Tanganyika Territory, south through Mozambique
(Amatongas) and the Rhodesias to Transvaal and Natal.

Gerrhosaurus nigrolineatus nigrolineatus Ilallowell
Black-lined Plated-Lizard
1857. Gerrhosaurus nigro-lineatus HaUowell, Proc. Acad. Nat. Sci. Phila-

"5 East African records of O. m. eechi Tornier ehoulcl be referred to wi. bottegoi,
as wi. zechi is the somewhat darker form ranging from the southern Sudan and
Belgian Congo west to Togo.

"G The extensive area of overlap with G. m. major is admitted and correct ; it
needs to be worked out in detail if m. grandis — apparently readily recognisable
in the south — is to be maintained.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 225

delphia, p. 49 : Gabon, French Congo.
1866. Gerrhosaurus invltilineatus Bocage, Jorn. Sci. Lisboa, 1. p. 44:

Duque de Braganca, Mossamedes, Angola.
1907. Gerrhosaurus flavigularis forma intermedia Lonnberg, in Sjostedt,
Wiss. Ergeb. Sc-hwed. Zool. Exped. Kilimanjaro, Meru umbeg.
Massaisteppen, No. 4, p. 7, pi. i, figs, la-lb: steppe near Lake
Natron, northern Tanganyika Territory.
1939. Gerrhosaunis nigrolin.eatus australis FitzSimons, Ann. Transvaal

Mus., 20, p. 10: Kaapmuiden, eastern Transvaal.
Range. Keiij'a Colony aud Tanganyika Territory, south to
the Transvaal, west to Southwest Africa, north to the French
Congo.

Gerrhosaurus flovigxilaris fitzsimonsi Loveridge
Kenya Yellow-throated Plated-Lizard
1942. Gerrhosaurus flavigularis fitzsimonsi Loveridge, Bull. Mus. Comp.

Zool., 89, p. 514: Mount Mbololo, Teita, Kenya Colony.
Range. Eastern Sudan and Ethiopia south through Kenya
Colony to southern Tanganyika Territory. ? "Zanzibar."

Genus TETRADACTYLUS Merrem

1820. Tetradactylu^ Merrem (not Dumeril & Bibron ms. Paris Mus.),
Vers. Syst. Amph., pp. 13, 75. Type by tautonomy: Chalcides
tetradactyhis Daudin.

1825. Cicigna Gray, Ann. Philos., (2) 10. p. 201. Type by monotypy:

Scincus Sep if or mis Schneider =z LaceHa seps Linnaeus.

1826. Saurophis Fitzinger, Neue Class. Eept., pp. 20, 50. Type by

monotjToy: Lacerta seps Linnaeus.
1838. Caitia Gray, Ann. Nat. Hist., 1. p. 389. Type by monotypy: C.

africana Gray.
1843. Eo-modactylus Fitzinger, Syst. Eept., p. 22. Type by original

designation: Caitia africana Gray.
1845. Pleurostrichus Gray, Cat. Lizards Brit. Mus., p. 51. Type by

present designation: Scincus seplfonnis Schneider = Lacerta

seps Linnaeus.
1892. Caetia Cope, Journ. Morph., 7, p. 224. Emendation for Caitia

Gray: 1838.

Tetradactylus fitzsimonsi simplex Laurent
Congo Long-tailed Seps
1950. Tetradactylus fitzsimonsi simplex Laurent, Revue Zool. Bot. Afr.,

43, p. 350: Kundelungu, 1750 metres, Belgian Congo.
Range. Southeastern Tanganyika Territory, west through
Northern Rhodesia to eastern Belgian Congo.

226 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Genus CHAMAESAURA Schneider

1801. Charnaesaura Schneider (part), Hist. Amphib., 2. pp. 205, 210.
Type by designation of Fitzinger :1826: Lacerta anguina Lin-
naeus.

1820. Monodaetylus Meirem, Vers. Syst. Amphib., pp. 13, 7G. Type by
mouotypy: Lacerta anguina Linnaeus.

1832. Cricochalcis Wiegmann, Handbueh Zool., p. 185. Type by mono-
typy : C. aenea Weigmann.

18G2. Muncvs Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 339. Type
by monotypy : M. maerolepis Cope.

Charnaesaura anguina tenuior Gunther Uganda Snake-Lizard
1895. Chaviacsaura tenuior Giinther, Ann. Mag. Nat. Hist., (6) 15, p. 524,
pi. xxi, fig. B : Kampala, Uganda.

1899. CUamu.esama anncctens Boulenger, Pi-oc. Zool. Soc. London, p.

97: Ravine Station, 7500 feet, Mau Mountains, Kenya Colony.
Range. Grasslands of eastern Belgian Congo, east through
Uganda and Kenya Colony to the Usambara Mountains, Tangan-
yika Territory.

Chamaesavura miopropus Boulenger
Northern Khodesia Snake-Lizard
1894. Charnaesaura miopropus Boulenger, Proc. Zool. Soc. London, pp.

724, 732: "Fwambo," i.e. Fwamba, Northern Ehodesia.
Range. Grasslands of southern Tanganyika Territory, west
through Northern Rhodesia to Angola.

Family LACERTIDAE"
Genus GASTROPHOLIS Pi.seher

1886. Gastropholis Fischer, Abhand. Nat. Ver. Hamburg, 9, p. 1. Type
by monotypy: G. vittata Fischer.

Gastropholis vittata Fischer Keel-bellied Ground-Lizard

1886. Gastropliolis vittata Fischer, Abhand. Nat. Ver. Hamburg, 9.
p. 1, pi. i, fig. 1: "Zanzibar," probably coast of opposite main-
land.

1900. Gastropholis lutsei Tornier, Zool. Jahrb. Syst., 13, p. 591, fig. — :

Tanga, Tanganyika Territory.

"7 lu his revision of this family, Boulenger (1920, Monog. Lacertitlae, 1, p. 2)
stressed the difliculties attending any attempt to arrange the genera in a linear
sequence, lie placed NucraK at the head of the list in the belief that it dated
from the Oligucene. This idea arose from a specimen preserved in wliat was
tliought to be Baltic amber, whereas it was almost certainly in so-calleil "Zanzi-
bar copal" from the Tanganyika coastlands.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 227

190-t. GastropJiolis prasiim Werner, Zool. Anz., 27. p. 462: Tanganyika
Territory (as Klceberg coll., probably Tanga).

Range. Coastal Tanganyika Territory and Mozambique (at
Lumbo. unless separable). No definite record from Zanzibar
Island.

Genus BEDRIAGAIA Boulenger

1916. Bedriagaia Boulenger, Ann. Mag. Nat. Hist., (8) 18. p. 112. Type
by monotypy: B. tiopidopholus Boulenger.

Bedriagaia moreaui Loveridge. Keel-bellied Tree-Lizard

193G. Bcdriagain moreaui Loveridge, Proc. Xew England Zool. Club, 15,
p. 67, figs. — : Ainani, Usambara Mountains, Tanganyika
Territory.

Range. Tanganyika Territory (a eanopy-dwelling species
known only from the holotype and a second specimen found in
tlie middle of Tanga town.sliip, so R. E. Moreau informs me —
6.ix.l944).

Genus HOLASPIS Gray

1863. HolaspL^ "A. Smith" Gray, Proc. Zool. Soc. London, p. 152. Tj-pe
by monotypy: E. guentheri Gray.

Holaspis guentheri laevis Werner
Eastern Serrate-toed Tree-Lizard
189.5. Holaspis guentheri laevis Werner, Verb. Zool. Bot. Ges. Wien, 45.
p. 191, pi. V, fig. 4: Usambara Mountains, Tanganyika Territory.
Range. Northeast Tanganyika Territory, south to Amatongas,
Mozambique.

Holaspis guentheri guentheri Gray

Western Serrate-toed Tree-Lizard
1863. Holaspis guentheri "A. Smith" Gray, Proc. Zool. Soc. London.

p. 153, pi. XX, fig. 1: No locality.
Range. TJganda, southwest to Angola, northwest to Sierra
Leone.

Genus PHILOCHORTUS Matschie

1893. Philochortus Matschie, Sitzb. Ges. Naturf. Freunde Berlin, p. 30.
Type by monotypy: P. neumanni Matschie.

228 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Philochortus intermedius rudoliensis Parker
Southern Shield-backed Ground-Lizard
1932. Philochortus intermedius rudolfcnsis Parker, Journ. Linn. Soc.
London, ZooL, 38, p. 226 : near mouth of Kaliokwell Eiver, Lake
Eudolf, Kenya Colony.
Range. Northern Kenya Colony (known only from the holo-
type).

Genus ALGYROIDES Bibron"

1833. Algyroides Bibron, in Bory de St. Vincent, Exped. Sci, Moree

Eept., p. 67. Type by monotyi^y: A. moreoticus Bibron.

1834. Notopholis Wiegmann (not Weigniann:1830), Herp. Mexicana, p.

10. Type by monotypy: N. fitsingeri Wiegmann.

1843. Tropidopholis Fitzinger, Syst. Eept., p. 21. New name for Noto-
pholis Wiegmann: 1834.

1868. Endromius Gistel, in Blicke, Leben Natur Menschen (Leipzig)
p. 150. Type {fide Mertens:1936) : Algyroides moreotictis
Bibron.

1912. Adolfus Sternfeld, Wiss. Ergebn. Deut. Zentral-Afrika-Exped.
1907-1908, 4. p. 220. Type by present designation; A. frideri-
cianus Sternfeld =: Algyroides afric<inus Boulenger.

Algyroides alleni Barbour Alpine-meadow Lizard

1914. Algiroides alleni Barbour, Proc. New England Zool. Club, 4. p. 97:

northeast slope of Mount Kenya, Kenya Colony.
Range. Al)ove 9000 feet on mountains of Uganda (Elgon) and

Kenya Colony (Aberdares; Kenya; Kinangop).

Algyroides africanus Boulenger Multi-scaled Forest-Lizard
1906. Algiroides africanus Boulenger, Proc. Zool. Soc. London, 2. p, 570,

fig. 96 : Entebbe, Uganda.
1912. Adolfus fridericianus Sternfeld, Wiss. Ergebn. Deut. Zentral-
Afrika-Exped. 1907-1908, 4, p. 220, figs. 1 and 2b: Avakubi.
upper Aruwimi Eiver, Belgian Congo.
Range. Uganda, west through the Belgian Congo to French
Cameroon.

"8 This name was proposed in a volume that appeared in 1833-1835. Sherborn
gives 1833 for Algyroides. but Boulensrer cites 1835, using the amended spelling
(Algiroides) proposed by Dumgril & Bibron, 1839, Erp^t. Gen. 5, pp. 174, 192.

Notopholis was first proposed by Wiegmann in 1830 to replace his preoccupied
Aplstes, of whicli the type was Lacerta edtcarsiana Dug6s = Psammosaurus
htapanicuB Fitzinger. Consequently Notopholis Wiegmann, 1834, is a preoccupied
honjonym.

LOVERIDQE: E. AFRICAN REPTILES AND AMPHIBIANS 229

Algyroides vcmereselli (Tornier) Sparser-scaled Forest-Lizard
1902. Lacerta vavcresclli Tornier, Zool. Anz., 25. p. 701: Kagera, west

of Lake Victoria, Tanganyika Territory.
1917. Algiroides Boul.engeri Peracca, Atti Accad. Sci. Torino, 52, p. 351 :

Fort Portal, Toro, western Uganda.
Range. Western Uganda and western Tanganyika Territory,
west through Belgian Ruanda-Urundi to the Belgian Congo.

Genus LACERTA Linnaeus

79

1758. *Lacerta Linnaeus (part), Syst. Nat. ed. 10, I, p. 200. Tj-pe by
designation of Fitzinger (1843:20): L. agilis Linnaeus.

1830. *Zootoca Wagler, Nat. Syst. Amphib., p. 155. Type by designation
of Fitzinger (1843:20): Lacerta crocea Wolf = L. vivipara
Jacques.

1830. *Podarms Wagler (part), Nat. Syst. Amphib., p. 155. Type by
designation of Fitzinger (1843:20): Seps muralis Laurenti.

1836. Timon Tscliudi, Isis von Oken, 29, p, 551. Type by original desig-
nation: Lacerta ocellata Daudin.

1838. Teira Gray, Ann. Nat. Hist., 1, p. 280. Type by monotypy: T.
punctata Gray ■= Lacerta dugesii Edwards.

1843. *Scelarcis Fitzinger, Syst. Kept., p. 20. Type by original designa-
tion: Lacerta perspicillata Dumeril & Bibron.

1843. J Zso(?rr)mw5 Fitzinger, Syst. Kept., p. 20. Type by original designa-
tion: Lacerta maderensis Fitzinger {nomen nudum).

1843. *Elaphropus Fitzinger, Syst. Kept., p. 20. Type by original designa-
tion: Lacerta galloti Dumeril & Bibron.

1843. Chrysolamprus Fitzinger, Syst. Kept., p. 20. Type by original
designation: Lacerta ocellata Daudin.

1843. Phenax Fitzinger, Syst. Kept., p. 20. Type by original designation
Lacerta taurica Pallas.

1845. Thetia Gray, Cat. Lizards Brit. Mus., p. 32. Type by monotypy:
Lacerta perspicillata Dumeril & Bibron.

1851. Atropis Gluckselig, Lotos, p. 138. Type by monotypy: Lacerta
nigra Wolf = a melanistic mutation of Lacerta vivipara Jacques.

1868. Agetosaura Gistel, in Blicke, Leben Natur Menschen (Leipzig),
p. 146. Type (fide Mertens:1936) : Lacerta margaritata Schinz.

1868. Tlialestris Gistel (not Glaus: 1863), in Blicke, Leben Natur

^ Six of these names Indicated by asterisks and here regarded as subgenera,
were treated as "Sections" by Boufenger (1920, Mono^r. Lacertidae, 1, p. 37)
though, as he ignored Fitzinger's names, he used Thetia (instead of Scelarcia)
and proposed Gallotia (instead of Elaphropus) . The only East African species,
L. jacksoni Boulenger, he refers to Podarcis, while vauereaelli Tornier (which he
had not seen) was assigned to Zootoca. I have since shown that it is an
Algyroidee, though with affinities to jackaoni that makes one wonder whether
they are not congeneric.

230 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Menschen (Leipzig), p. 148. Type {fide Mertens:1936) : Lacerta

nigi-a Wolf.
18G8. Tritonopsis Gistel (not Conrad .-1865), in Blicke, Leben Natur

Menschen (Leipzig), p. 148. Type (fide Mertens:1936), the

same as for Thalestris.
1916. Gallotia Boulenger, Trans. Zool. Soc. London, 21, p. 3. Type by

tautonomy : Lacerta galloti Dumeril & Bibron.
1916. *Centromastix Boulenger, Trans. Zool. Soc. London, 21. p. 3. Type

by monotypy : Lacerta echinata Cope.

Lacerta jacksoni Boulenger JacLson's Lizard

1899. Lacerta jaclcsoni Boulenger, Proc. Zool. Soc. London, p. 96, pi. x:

Ravine Station, Mau Mountains, Kenya Colony.
1907. Lacerta jaclcsoni hibonote7iffis Lonnberg, in Sjostedt, Wiss. Ergeb.

Schwed. Zool. Exped. Kilimanjaro, Meru umbeg. Masaaisteppen,

No. 4, p. 5: "Kibonoto," i.e. Kibongoto, Kilimanjaro, Tan

ganyika Territory.
Range. Uganda; Kenya Colony and Tanganyika Territory,
west through Belgian Riianda-Urundi to the Belgian Congo.

Genus NUCHAS Gray

1838. Xucrafi Gray, Ann. Nat. Hist., 1. p. 280. Type by monotypy:
Lacerta lalandii Milne-Edwards.

1843. Heliophilus Fitzinger, Syst. Eept., p. 20. Type by original designa-
tion: Lacerta delalandii (sic) Milne-Edwards.

1886. Bettaia Bedriaga, Abhand. Senckenberg. Ges., 14, No. 2, p. 24.
Type by monotypy: Lacerta delalandii (sic) Milne-Edwards.

Nucras boulengeri boulenger! Neumann*"
Uganda Savanna-Lizard

1900. Nucras boulengeri O. Neumann, Ann. Mag. Nat. Hist., (7) 5,

p. 56: "Lubwa's, " i.e. Luba's Village, Busoga, Uganda.
1907. Nucras emini Boulenger, Ann. Mag. Nat. Hist., (7) 19, p. 488:

southern shore of Lake Victoria, Tanganyika Territory.
1909. Nucras ukerewensis Bolkay, Archivum Zool. Budapest, 1, p. 13,

so If , as I believe, Nucras succlnea Boulenger (1917, Ann. S. Africa Mus., 13,
p. 195, footnote) was in Zanzibar copal (instead of Baltic amber as alleged),
then that name should be added to the synonymy. However, the status of these
synonyms needs claritication, for two color forms appear to be involved — the
slender, rufous lizard frequenting laterite areas of Tanganyika where it is to be
found on native paths, and a more robust, dark olive, multiocellate, northeastern
race represented in the Museum of Comparative Zoology by a series from Lolta
Plains (M.C.Z. 17985-9) to which the name ukerewensis Bolkay seems applicable.
Why Bolkay's single subadult ^ holotype should have been so named is not
clear. I saw no sign of the species during my short stay on Ukerewe Island in
19.30.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 231

figs. 1-4: Shirati, east shore of Lake Victoria, Tanganyika
Territory.
Range. Uganda (east bank of Nile), east to Kenya Colony
(Eldama Ravine), south through Tanganyika Territory (except
around Kilosa) to the Rovuma River.

Nucras boulengeri kilosae Loveridge Keeled Savanna-Lizard
1922. Nucras kilosae Loveridge, Proc. Zool. Soc. London, p. 314: KUosa,

Usagara, Tanganyika Territory.
Range. Country around Kilosa, east-central Tanganyika Ter-
ritory.

Genus LATASTIA Bedriaga

1884. Latastia Bedriaga, Ann. Mus. Civ. Stor. Nat. Genova, 20, p. 307.

Type by present designation: Lacerta samhnrica Blanford ■=■ L.

longicandata Reuss.
1907. Apdihya Mehely, Potfiiz. Termesz. Koslony (Budapest), 85, p. 26.

Type by monotypy: Lacerta cappadocdca Werner.

Latastia longicaudata revoili (Vaillant)^^
Southern Long-tailed Lizard
1882. Eremias revoili YaOlant, Miss. Revoil Pays Qomal., Rept., p. 20,

pi. iii, fig. 2 : Somaliland.
1884. Latastia doriai Bedriaga, Ann. Civ. Stor. Nat. Genova, 20, p. 313 :

"Rugdeia Sogheira (Adal)."
1884. ILatastia doriai var. tnartensi Bedriaga, Ann. Mus. Civ. Stor. Nat.

Genova, 20, p. 316: Keren, Bogos, Eritrea.
Range. (Northern limits uncertain; apparently arid areas of)
Eritrea, south through Ethiopia and Kenya Colony to Ugogo,
central Tanganvika Territory.

Latastia johnstoni Boulenger Nyasaland Long-tailed Lizard
1907. Latastia johnstoni Boulenger, Ann. Mag. Nat. Hist., (7) 19, p.

392: "Masuka, " i.e. Masuku, and Nyika Plateaus, Nyasaland.®-
1919. Latastia TcidwelU Boulenger, Trans. Royal Soc. S. Africa, 5, p. 39:

Eldorado, Southern Rhodesia.
1942. Latastia hredoi de Witte, Bull. Mus. Royal Hist. Nat. Belgique,

18, p. 1, figs. 1-2: Musosa, Tanganyika District, Belgian Congo.

81 Z/. doriai appears to be intermediate between L. I. longicaudata (Reuss)
and revoili (Vaillant) ; whether it is subspecifically recognizable is somewhat
doubtful though Parker (1942, Bull. Mus. Comp. Zool., 91, p. 69) inclines to
think it may prove so. Geographically its position is far from clear and calls
for more material from the general region.

S2 Possibly from the lower slopes under 4000 feet, not from the plateaus (cf.
Loveridge, 1953. BuU. Mus. Comp. Zool., 110, p. 229).

232 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Range. Central Tanganyika Territory, south through Mozam-
bique; Nyasaland and both Rhodesias to the eastern Belgian
Congo.

Genus EREMIAS Wiegmann^

1834. *Eremias Wiegmann, Herp. Mexicana, p. 9. Type by subsequent
designation of Boulenger:1918: Lacerta velox Pallas.

1838. *Mesalina Gray, Ann. Nat. Hist., 1. p. 282. Type by monotypy:
M. lichtensteinii Gray = Lacerta rubropunctata Lichtenstein.

1841. Aspidorhinus Eichwald, Fauna Caspio-Caucasia, p. 74. Type by
monotypy: A. gracilis Eichwald = Lacerta velox Pallas.

1843. Eremioscopiis Fitzinger, Syst. Eept., p. 20. Type by original desig-
nation : Lacerta guttulata Lichtenstein.

1843. *neUobolus Fitzinger, Syst. Rept., p. 20. Type by original designa-
tion: Lacerta lugubris A. Smith.

1843. Fataeosaurus Fitzinger, Syst. Eept., p. 20. Type by original desig-
nation: Lacerta capensis A. Smith (not Sparrman:1783).

1843. Fediophylax Fitzinger, Syst. Kept., p. 21. Type by original desig-
nation: Eremias dorsalis Dum6ril & Bibron =: Lacerta lugubris
A. Smith.

1843. Choroscopus Fitzinger, Syst. Eept., p. 21. Type by original desig-
nation: Eremias Uneo-ocellata Dumeril & Bibron.

1843. Eudioptra Fitzinger, Syst. Eept., p. 21, Type by original designa-
tion : Lacerta undata A. Smith.

1843. Dioptroblepharis Fitzinger, Syst. Eept., p. 21. Type by original
designation : Eremias pardalis Dumeril & Bibron (not of Lich-
tenstein) =: Lacerta guttulata Lichtenstein.

1843. Pedioplanis Fitzinger, Syst. Eept., p. 21. Type by original designa-
tion : Eremias burchelli Dumeril & Bibron.

1843. Tachyscelis Fitzinger, Syst. Eept., p. 21. Type by original designa-
tion: Lacerta rubropjinctata Lichtenstein.

1883. *Pseuderemias Boettger, Abhand. Senckenberg. Naturf. Ges. 13.
p. 118, footnote. Type by monotypy: P. lineolata "Euppell"
Boettger = Acanthodactylus mucronatus Blanford.

1885. Boulengeria Lataste, Ann. Mus. Civ. Stor. Nat. Genova, (2) 2,
p. 126. Type by subsequent designation of Boulenger (1918) :
AcuntTiodactylus mucronatus Blanford.

1918. Lampreremias Boulenger, Journ. Zool. Eos., 3. p. 2. Type by
original designation: Eremias nitida Giinther.

S3 pive of these names, indicated by asterisks, correspond to "Sections" in
Boulenger (1921, Monogr. Lacertitlae, 2, pp. 227-231) though his Lampreremias
is antedated by Heliobolua Fitzinger, whose names he ignored — and not without
reason for seven of Fitzinger's names are synonymous with Mesalina Gray.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 233

1918. *'Taniereinias Boulenger, Journ. Zool. Res., 3, p. 2. Type by original

designation: Eremias guineensis Boulenger,
1928. Ommateremias Lantz, Bull. Muzei Gruzii, Tiflis, 4, p. 37. Type

by original designation: Lacerta arguta Pallas.

Eremias neumanni Tornier Neumann's Sand-Lizard

1905. Eremias neumanni Tornier, Zool. Jahrb. Syst., 22. p. 376:
"Barssa" i.e. Brussa Valley, north of Lake Stephanie, Ethiopia.
Range. Ethiopia, south to Ngatana, Tana River, Kenya Col-
ony.

Ermias spekii sextaeniota Stejneger
Northern Speke'.s Sand-Lizard
1893. Eremias sextaeniata Stejneger, Proc. U. S, Nat. Mus., 16. p. 718:

Tana Eiver, Kenya Colony,
Range. Southern Sudan and northern Uganda, east to British
Somaliland, south through Somalia to northern Kenya Colony.

Eremias spekii spekii Giinther Southern Speke's Sand-Lizard
1872. Eremias spehii Giinther, Ann. Mag. Nat, Hist., (4) 9. p. 381:

Unyamwezi, Tanganyika Territory.
1878. Eremias riigiceps Peters, Monatsb. Akad, Wiss, Berlin, p. 202, pi,

ii, fig. 1: "Taita," i.e. Teita, Kenya Colony.
Range. Kenya Colony (south of Tana River), south to central
Tanganyika Territory.

Eremias smithi Boulenger Smith's Sand-Lizard

1895. Eremias smithi Boulenger, Proc. Zool, Soc, London, p. 534, pi.

xxix, fig. 4: Milmil, in Haud, Ethiopia.
Range. Ethiopia and British Somaliland, south through

Somalia to the Tana River, Kenya Colony.

Eremias striata Peters Peter's Sand-Lizard

1874. Eremias hrenneri var. striatvs Peters, Monatsb. Akad. Wiss. Ber-
lin, p. 370: "Barawa, " i.e. Brava, Somalia.
1893. Eremias hoehneli Stejneger, Proc. U, S. Nat. Mus., 16. p, 719:

Tana Eiver, Kenya Colony,
Range. British Somaliland, south through Somalia to the
Tana River, Kenya Colony.

Genus ICHNOTROPIS Peters

1843. Thermophilus Fitzinger (not Thermophila Hubner:1816), Syst.
Rept., p, 21, Type by original designation: Tropidosaura capen-
sis Dumeril & Bibron = Algyra capensis A. Smith.

234 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1854. Ichnotropis Peters, Monatsb. Akad. Wiss. Berlin, p. 617. Type by
subsequent designation: 7. macrolepidota Peters = Algyra
capensis A. Smith.

Ichnotropis tanganicana Bouleuger

Tanganyika Rough-scaled Sand-Lizard
1917. Ichnotropis tanganicana Boulenger, Ann. Mag. Nat. Hist., (8) 19,
p. 278: East coast of Lake Tanganyika, Tanganyika Territory.
Range. Tanganyika Territory (known only from the holo-
type).

Ichnotropis capensis bivittata Bocage
Angola Rough-scaled Sand-Lizard
1866. Ichnotropis bivittata Bocage, Jorn. Sci. Lisboa, 1, p. 43: Duque

de Braganea, Angola.
Range. Southern Tanganyika Territory, west through Belgian
Congo to Angola, possibly north to French Congo.

Ichnotropis squamulosa Peters

Mozambique Rough-scaled Sand-Lizard
1854. Ichnotropis squamulosa Peters, Monatsb. Akad. Wiss. Berlin, p.

617: Tete, Mozambique.
Range. Tanganyika Territory, south through Mozambique to
Zululand, Natal, west through Transvaal and Bechuanaland to
Southwest Africa, north to Angola.

Family VARANTDAE
Genus VARANUS Merrem'*

1818. Monitor Liohtenstein (not Blainville:1816), Zool. Mus. Univ. Ber-
lin (2), p. 66. Type by tautonomy: Lacerta monitor Linnaeus
(part) = L. nilotica Linnaeus (part).

1820. *Varanus Merrem, Vers. Syst. Amphib., pp. 13, 58. Type by sub-
sequent designation: Laceria varia Shaw.

1826. *Psammosaurus Fitzinger, Neue Class. Kept., pp. 21, 50. Type by

monotypy: Tupmambis griseu^ Daudiu.

1827. Uaraiuis Gray, Philos. Mag., (n.s.) 2. p. 55. Emendation of

Varanus Merrem.
1827. Draca,e7M Gray (not Daudiu:1802), Philos. Mag., (n.s.) 2. p. 55.

8-t Eight of these names, indicated by asterisks, are valid subgenera according:
to Mertens (1942, Abhaud. Senckeuberg. Naturf. Ges., No. 4(56, p. 242: key)
whose comprehensive and careful revision of the VARANIDAB has been of
the greatest help.

L0\T:RIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 235

Type by tautonomy: Lacerta dracnena Linnaeus = TupinamMs
bcngalensi^ Daudin.
1830. Hydrosaurus Wagler (not Kaup:1828), Nat. Syst. Amphib., pp.

132, 164. Type by subsequent designation: Tnpinambis bivit-

tatus Kuhl ^ Stellio sahator Laurenti.
1830. *PoUjdaedalm Wagler, Nat. Syst. Amphib., pp. 132, 164. Type by

subsequent designation: Lacerta nilotica Linnaeus.
1838. *Empagusia Gray, Ann. Nat. Hist., 1. p. 393. Type by subsequent

designation: Manitor flavescens Hardwicke & Gray.
1838. *Odatria Gray, Ann. Nat. Hist., I. p. 394. Type by monotypy: 0.

punctata Gray = Monitor tristis Schlegel.
1843. Cylindruriis Fitzinger, Syst. Eept., p. 19. Type by original desig-
nation. Odatria punctata Gray = Monitor tristis Schlegel.
1843. Panther osaur us Fitzinger, Syst. Eept., p. 19. Type by original

designation: Eydrosaurus gouldii Gray.
1843. Acjalynatosaurus Fitzinger, Syst. Kept., p. 19. Type by original

designation: Monitor timorensis Gray.
1843. Euprepiosaurus Fitzinger, Syst. Eept., p. 19. Type by original

designation: Varanu^ chlorostigma Dumeril & Bibron = Tupi-

nambis indicus Daudin.
1843. Pachysauru-s Fitzinger, Syst. Eept., p. 20. Type by original

designation: Tupinambis albigularis Daudin.
1843. Bhinoptyon Fitzinger, Syst. Eept., p. 20. Type by original designa-
tion: Varanus ocellatus Eiippell = Lacerta exanthemo,tic<i Bosc.
1843. Psammoscopus Fitzinger, Syst. Eept., p. 20. Type by original

designation: Varanus picquotii Dumeril & Bibron = Monitor

flavescens Hardwicke & Gray.
1845. Eegenia Gray, Cat. Lizards Brit. Mus., p. 8. Type by subsequent

designation: Tupinambis albigularis Daudin.
1927. Placovaranus Fejervary, Termeszettudomanyi Kozlony, 59. p. 284.

Type by original designation: Varanus Tcomodoensis Ouwens.
1942. *Indovaranus Mertens, Abhand. Senckenberg. Naturf. Ges., No.

466, p. 240. Type by original designation: Tupinambis benga-

len-sis Daudin.
1942. *Dendrovaranus Mertens, Abhand. Senckenberg. Naturf. Ges., No,

466, p. 241. Type by original designation: Varanus rudicollis

Gray.
1942. *Tectovaranus Mertens, Abhand. Senckenberg. Naturf. Ges., No.

466, p. 242. Type by original designation: Monitor dumerili

Schlegel.

Varanus niloticus niloticus (Linnaeus) Nile Monitor

1758. Lacerta monitor Linnaeus (part), Syst. Nat., ed. 10, 1. p. 201:
"in Indiis, " i.e. Africa.

236 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1766. Lacerta nilotica Linnaeus, Syst. Nat., ed. 12, 1, p. 369: Egypt.

1768. Stellio saurus Laurenti, Syn. Eept., i>. 56: "in Zeylania ad littora
maris. ' '

1783. Lacerta capensis Sparrman, Eesa till Goda Hoppsudden, p. 749:
" Hinterbruyntjes Heights," = Bruintjeshoogte, a village be-
tween Pearston and Somerset East, eastern Cape Province.^^

1788. " L. tupinamhis" LacepMe (part),*^ Hist. Nat. Quad. ovip.
Serpens, 1, Synopsis Methodica, p. 251, pi. xvii: restricted to
Cape of Good Hope, i.e. South Africa.

1802. Tupinavihis elegans Daudin (part). Hist. Nat. Rept., 3, p. 36:
"Amerique meridionale" (error).

1802. Tupinamhis stellatus Daudin, Hist. Nat. Eept., 3, p. 59, pi. xxxi:
Senegal (restricted).

1819. Monitor pultiher Leach, in Bowdich, Miss. Ashantee, App., p. 493:
"Fantee, " i.e. Fanti, Ashanti, Gold Coast.

1844. Monitor elegans senegalensis Schlegel, Abbild. Amphib., p. x:

Senegal.
Range. Africa (exclusive of arid areas in the north and south-
west, also the rain-forest region of the west^'), including Uganda;
Kenya Colony ; Tanganyika Territory ; Zanzibar and Mafia
Islands.

Voranus exanthematicus microstictus Boettger''*
Eastern Savanna-Monitor

1845. Mo>nitoT microstictus Riippell {nomen nudum), Mus. Senckenberg.,

3. p. 301: " Abyssinien, " i.e. Ethiopia.
1893. Varanus microstictus Boettger, Kat. E^pt. Saminl. Mus. Sencken-
berg. Naturf. Ges., Part 1, pp. viii, 72: Ethiopia.
Range. Ethiopia and British Somaliland, south through
Uganda ; Kenya Colony ; Tanganyika Territory and Zanzibar
Island to Mozambique (where it meets with the southern race,
V. e. alhigularis (Daudin)).

ȣ- Dr. V. F. FitzSimons, to whom I am indebted for the above information,
informs me that this locality appears on Sparrman's map (1876) as "Agter-
bruntjes Hoogte."

86 Only tlie vernacular name is given, not "Lacertus tupinamhis" as cited by
some authors. In fact it is listed with the crocodiliaus in the Synopsis
Methodica.

87 The only race, V. n. ornatus (Daudin), has a discontinuous distribution in
the surviving patches of rain forest from the western Belgian Congo and
Cameroons to Liberia. The species is referable to the subgenus Polydaedalus.

88 This is the monitor which for a century has been miscalled Varanus ocellatus
(Heyden in Kuppell), a name that is actually synonymous with V. e. exanthe-
maticus (Bosc) which ranges from Eritrea to Senegal.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 237

Family AMPHISBAENIDAE
Genus ANCYCLOCRANIUM Parker

1942. Ancyclocranium Parker, Bull. Mus. Comp. Zool., 91, p. 37: Type
by original designation: Anops somalicus Scortecci.

Ancyclocronimn borkeri Loveridge

Lindi Sharp-snouted Worm-Lizard
1942. Ancyclocranium iarlceri Loveridge, Proc. Biol. Soc. Washington,
59. p. 73, pi. xiii: Mbemkuru Eiver, Lindi district, Tanganyika
Territory.
Range. Tanganyika Territory (known only from holotype).

Ancyclocranium ionidesi Loveridge
Kilwa Sharp-snouted Worm-Lizard
1955. Ancylocranium ionidesi Loveridge, Journ. E. Africa Nat. Hist.
Soc, 22, p. 177, pi. — : Kilwa, Southern Province, Tanganyika
Territory.

Range. Tanganyika Territory (known only from the type
series).

Genus AMPHISBAENA Linnaeus *='

1758. * AmpMsljaena Linnaeus, Syst. Nat. ed. 10, 1, p. 229. Type by sub-
sequent designation: A. fidiginosa Linnaeus.
1844. Sarca Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 71. Type by

monotypy: Amphisbaena caeca Cuvier.
1844. *Cynisca Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 71. Type

by monotypy : Amphisbaena leucura Dumeril & Bibron.
1861. *Diphaluti Cope, Proc. Acad. Nat. Sei. Philadelphia, p. 75. Type

by original designation : D. fenestratua Cope.
1865. *Bronia Gray, Proc. Zool. Soc. London, p. 448. Type by monotypy:

B. brasillana Gray.
1878. Ophloproctes Boulenger, Bull. Soc. Zool. France, 3, p. 300. Type

by monotypy : 0. liberiensis Boulenger.
1885. Aporarclius Cope, Proc. American Philos. Soc, 22, p. 187. Type

by monotypy: Amphisbaena prunicolor Cope.
1885. Zygnis Cope (not Oken:1816), Proc. American Philos. Soc, 22, p.

188. Type by monotypy: Amphi-sbaen-a quadrifrons Peters.
1907. *Cluriiulia Boulenger, Ann. Mag. Nat. Hist., (7) 20, p. 48. Type

by monotypy: C. swynnertoni Boulenger.

89 Seven of these names, indicated by asterisks, here regarded as subgenera, are
considered full genera by Vauzolini (1951, Herpetologica. 7, pp. 113-123). Five of
the six East African species would be assigned to as many different subgenera,
with his Hhrevea replacing Zygnis Cope (preoccupied).

238 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1911. Amphisbaeniila Sternfcld, Sitzb. Ges. Naturf. Fieunde Berlin,
p. 246. Type by monotypy: A. orientalis Sternfeld.

1951. *Loveridgea Vanzolini, Herpetologica, 7. p. 114. Type by original
designation: Aviphishaena phylofiniens Tornier.

1951. *Shrevea Vanzolini, Herpetologica, 7. p. 115. Type by original
designation: Amphisbaena quadrifrons Peters.

Amphisbaena ionidesii Battersby Liwale Worm-Lizard

1950. Ampliishaena ionidesii Battersby, Ann. Mag. Nat. Hist., (12) 3,
p. 413, fig. — : Liwale, Southern Province, Tanganyika Terri-
tory.
Range. Tanganyika Territory.

Amphisbaena phylofiniens Tornier Ujiji Worm-Lizard

1899. Amphishaoui phylofiniens Tornier, Zool. Anz., 22. p. 260: Ujiji,

Tanganyika Territory.
Range. Tanganyika Territory.''"

Amphisbaena mpwapwaensis Loveridge
Mpwapwa Worm-Lizard

1932. Amphisbaena mpwapwaensis Loveridge, BuU. Mus. Comp. Zool.,

72. p. 378: Mpwapwa, Ugogo, Tanganyika Territory.
Range. Tanganyika Territory.^"

Amphisbaena ewerbecki (AVeruer) Mbanja Worm-Lizard

1910. Chirindia ewerbecM Werner, 1909, Jalirb. Hamburg, Wiss. Anst.,

27, p. 37: Mbanja near Lindi, Tanganyika Territory.
Range. Tanganyika Territory.^"

Amphisbaena rondoensis Loveridge Nchingidi Worm-Lizard
1941. Amphishaena rondoensis Loveridge, Bull. Mus. Comp. Zool., 87,
p. 394, fig. 23: Nchingidi, Rondo Plateau, Lindi district, Tan-
ganyika Territory.

Range. Tanganyika Territory.®**

Amphisbaena orientalis (Sternfeld) Mikindani Worm-Lizard

1911. Amphisbaenula orientalis Sternfeld, Sitzb. Ges. Naturf. Freunde

Berlin, p. 246: Mikindani, Southern Province, Tanganyika Ter-
ritory.
Range. Tanganyika Territory .°"

Genus GEOCALAMUS Giinther

1880. Geocalamus Giinther, Ann. Mag. Nat. Hist., (5) 6. p. 234. Type
by monotypy: G. modestiis Giinther.

90 Only from ^icinity of the t.vpe localit.v.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 239

Geocolomus acutus Sternfeld

\"oi Wedg'e-siKJiited AVorm-Lizard

1912. CTeocalamii.s acutus Sternfeld, Wiss. Ergebn. Deut. Zentral-Afrika-

Exped. 1907-1908, 4. p. 209: Voi. Kenya Colony.

1913. deocalamus noltci Boettger, in Voeltzkow, Eeise in Ostafrika, 3,

p. 366, pi. xxvi, tig. 6: Moslii, Tanganyika Territory.
Range. Kenya Colony and Tanganyika Terirtory.

Geocalamus modestus Giinther

]\I])^vap^va Wedge-.snouted Worm-Lizard
1880. Geocalamus modestvs Giinther, Ann. Mag. Nat. Hist., (5) 6. p.

23-1: Mpwapwa, Ugogo, Tanganyika Territory.
Range. Tanganyika Territory.

Suborder SERPENTES ''
Family TYPHLOPIDAE
Genus TYPHLOPS Oppel"

1811. Tijphlops Oppol, Ordn. Fam. Gattung Eept., p. 54. Type by sub-
sequent designation: Anguis lumbricalis Linnaeus.

1830. Typhlin-a Wagler (not TypJtline Wiegmann), Xat. Syst. Amphib.,
p. 196. Type by subsequent designation: " Acontias lineatiis
Eeinw. " = Typhlops Uneatus Boie.®"

1843. AspUlorln/nclius Fitzinger, Syst. Rept., p. 24. Type by original
designation: Typhlops eschrichtii Schelegel ^ Acontias pioictatus
Leach.

1843. Gerrhopilus Fitzinger, Syst. Eept., p. 24. Type by original designa-
tion: Typhlops ater Schlegel.

1843. Pseudotyphlops Fitzinger, Syst. Eept., p. 24. Type by original
designation : Typhlops polygrammicus Schlegel.

1843. Rhinotyphlops Fitzinger, Syst. Eept., p. 24. Type by original
designation: Typhlops lahindli Schlcgc-l.

1843. Eamphotyphlops Fitzinger, Syst. Eept., p. 24. Type by original
designation: Typhlops multilineatits Schlegel.

910PHIDIA of Macartney (1802), Boulengcr (1893) and others.

93 In case Fitzinger's (1843) designation or later ones be considered inadequate,
the type of TyphUna is hereby fixed as above to avoid any possibility of Typhlops
Keptemstrintus Schneider being selected, for this would affect the use of Lepto-
tuphlops and LEPTOTYPHLOPIDAE.

240 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

1844. Pilidion Diimeril & Bibron, Erpet. Gen., 6, p. 257. New name for
Typhliiia lineatus Wagler ^ TypMops lineatus Boie.

1844. Ophthalmidium Dumeril & Bibron, Erp6t. G6n., 6. p. 262. Type
by subsequent designation: 0. longissimum Dumeril & Bibron.

1844. Cathetarhinus Dumeril & Bibron, Erp6t. Gen., 6. p. 268. Type by
monotypy: C. melanoeephalus Dumeril & Bibron.

1844. Onychocephalus Dumeril & Bibron, Erpet. Gen., 6. p. 272. Type

by subsequent designation: TypMops lalandii Sclilegel.

1845. Onychophis Gray, Cat. Lizards Brit. Mus., p. 132. Type by

monotypy: 0. franMinii Gray = TypMops lalandii Schlegel.
1845. TypMinalis Gray, Cat. Lizards Brit. Mus., p. 134. New name for

TypMina Wagler.
1845. Anilios Gray, Cat. Lizards Brit. Mus., p. 135. Type by present

designation: A. Icachii Gray = Anguis lumbricalis Linnaeus.
1845. Argyrophis Gray, Cat. Lizards Brit. Mus., p. 136. Type by present

designation: A. bicolor Gray = TypMops nigroalbiis Dumeril &

Bibron.
1845. Meditoria Gray, Cat. Lizards Brit. Mus., p. 139. Type by mono-
typy: M. nasuta Gray = Anguis lumbricalis Linnaeus.
1861. DiaphorotypMops Jan, Arch. Zool. Anat. Fisiol., 1, p. 185. Type

by subsequent designation: D. disparilis Jan.
1868. Letheobia Cope, Proe. Acad. Nat. Sci. Philadelphia, p. 322. Type

by designation of Dunn & Dunn (1940: Copeia, p. 75): L.

pallida Cope.
1881. G-rypotypMops Peters, Sitzb. Ges. Naturf. Freunde Berlin, p. 70.

Type by original designation: Onydhocephalus acutus Dumeril

& Bibron.

Typhlops blanfordii bloniordii Boulenger
Ethiopian Blind-Snake
1889. Typhlops Blanfordii Boulenger, Ann. Mag. Nat. Hist., (6) 4.

p. 363: Senafe, Ethiopia.
Range. Eritrea and Ethiopia, south to northern Kenya Col-
ony (at Moyale; fide Scortecei:1940).

Typhlops blanfordii lestradei Witte^ Ruanda Blind-Snake
1933. Typhlops Lestradei de Witte, Revue Zool. Bot. Afr., 23. p. 206,
figs. 1-3: "Rubengera" i.e. Ruhengeri, Belgian Ruanda-Urundi.
Range. Uganda, west through Belgian Ruanda-Urundi to

Belgian Congo.

Typhlops schlegelii brevis Scortecci
Northern Schlegel's Blind-Snake

9* For detailed discussion of relationsliips with adolfi Sternfeld and dtihius
Cliabanaud, sec Loveridge, 1942, Bull. Mus. Comp. Zool., 91, pp. 254-255, pi. 11,
fig. 2.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 241

1929. Typhlops brevis Scortecci, Atti Soc. Ital. Sci. Nat. (Milano), 68,
p. 287: Chisiniaio (Kismayu), Somalia.

Range. Southern Sudan and Uganda (Lado Enclave) ; British
Somaliland south through Somalia (probably to northern Kenya
Colony).

Typhlops schlegelii excentricus Procter
Dark-bellied Blind-Snake
1922. Typhlops excentricus Procter, Ann. Mag. Nat. Hist., (9) 9, p. 685:

Kilosa, Tanganyika Territory.
Range. Tanganyika Territory (an east-central area from
Mpwapwa to Morogoro).

Typhlops schlegelii mucruso (Peters)®'^
Eastern Schlegel's Blind-Snake
1854. Onychoccphalus mucruso Peters (part), Monatsb. Akad. Wiss.

Berlin, p. 621 : Maeanga, Mozambique.
1873. Onychocephahis Pctersii Bocage, Jorn. Sci. Lisboa, 4. p. 249:

Bibala, Mossamedes, Angola.
1886. Typhlops {Onychoccphalus) hnmbo Bocage, Jem. Sci. Lisboa, 11-

p. 171 : Quisange, Benguela, Angola.
1893. Typhlops hottentotus Bocage, Jorn. Sci. Lisboa, (2) 3. p. 117:

Quindumbo, Benguela, Angola.
1893. Typhlops mandensis Stejneger, Proc. U. S. Nat. Mus., 16, p. 725:

Wange, mainland opposite Manda Island, Kenya Colony.
1910. Typhlops latirostris Stemfeld, Mitt. Zool. Mus. Berlin, 5. p. 70:

Tabora, Tanganyika Territory.
1912. Typhlops viridiflavus Peracca, Ann. Mus. Zool. Univ. Napoli, (2)

3, No. 25, p. 3 : Lake Bangweulu, Northern Rhodesia.
Range. Kenya Colony; Tanganyika Territory and Zanzibar
Island, south to Mozambique north of the Zambezi River,^'^ west
through Nyasaland and Northern Rhodesia to Angola.

Typhlops punctalus punctatus (Leach)®' Spotted Blind-Snake

1S19. Acontias punctatus Leach, in Bowdich, Mis.s. Ashantec, App., p.

493: "Fantee," i.e. Fanti, Ashanti, Gold Coast.
1844. Typhlops cschrichiii Schlegel, Abbild. Amphib., p. 37, pi. xxxii,

figs. 13, 16: Gold Coast.

^■' Owing to young individuals having rounded snouts resembling those of
})unctatus, they have often been reported as that species. Direct comparison oC
the types of the numerous synonyms of both species is very mucli uceded.

"•5 An arbitrary boundary, for the two races merge in this vicinity. T. s.
schlegelii (Peters) ranges over the whole of Africa south of the river.

9" Several other forms, which should porhaps be added to this synonymy, have
been described in the last thirty-live years.

242 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1844. Onydhocephalus congestus Dum^ril & Bibron, Erpet. Gen., 6. p.

265 : No locality.

1845. OnychopJiis Barroivii Gray, Cat. Lizards Brit. Mus., p. 133:

"India?"
1848. Onycltoccphalus Liheriensis Hallowell, Proc. Acad. Nat. Sci. Phila-
delphia, p. 59: Liberia.
1848. Onycliocephalus niciro-lineatus Hallowell, Proc. Acad. Nat. Sci.

Philadelphia, p. 60 : Liberia.
1864. Typhlops liheriensis var. intermedia Jan, Icon. Gen. Ophid., p. 24,

and 1, livr. 5, pis. v-vi, fig. 2: Liberia.
1864. Typhlops Kraussi Jan, Icon. Gen. Opliid., p. 26, and 1, livr. 3,

pi. vi, tig. 2 : Gold Coast.
1864. TypMops lineolatus Jan. Icon. Gen. Ophid., p. 24, and 1. livr. 9,

pi. i, fig. 4 : Sierra Leone.
1866. Onycliocephalus angolensis Bocage, Jorn. Sci. Lisboa, 1, p. 46:

Duque de Braganca, Angola.
1887. Typhlops (Onychocephalus) congicus Boettger, Zool. Anz., 10, p.

650: Povo Netonna, near Banana, Belgian Congo.
1893. Typhlops houlengcri Bocage, Jorn. Sci. Lisboa, (2) 3. p. 117:

Interior of Benguela, Angola.
1904. Typhlops iocagei, Perreira, Jorn. Sci. Lisboa, (2) 7, p. 114:

Cabicula, Bom Jesus, Angola.
1910. Typhlops Tornieri Sternfeld, Mitt. Zool. Mus. Berlin, 5. p. 69:

Mount Kilimanjaro, Tanganyika Territory.
1910. Typhlops Adolphi Sternfeld, Mitt. Zool. Mus. Berlin, 5. p. 70:

"Fort Blus," later corrected to Port Beni, Belgian Congo.
1917. Typhlops dtibiiis Chabanaud, Bull. Mus. Hist. Nat. (Paris), 1916,

22. p. 364: "Kivori, " i.e. Kivu Volcanoes, 1500 metres, Belgian

Congo.
1920. Typhlops Milleti Chabanaud, Bull. Mus. Hist. Nat. (Paris), 26.

p. 463: Togo.
Range. Forested areas of Uganda; montane forests of Kenya
Colony and Tanganyika Territory, southwest to Angola and
northwest to Senegal (frequently recorded in error for other
species).

Typhlops punctatus gierrcd Mocquard
Usambara Spotted Blind-Snake
1897. Typhlops Gierrai Mocquard, Bull. Mus. Hist. Nat. (Paris), 3.

p. 122: Tanga, Tanganyika Territory.
Range. Virgin forests of the Usambara and Uluguru Moun-
tains (possibly the gallery forests of adjacent lowlands also),
Tanganyika Territory.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 248

Typhlops kaimosae Loveridge Kakamega Spotted Blind-Snake
1935. Typhlops kaimosae Loveridge, Bull. Mus. Comp. Zool., 79, p. 5:

Kaimosi, Kakamega, Nyanza Province, Kenya Colony.
Range. Kenya Colony (known only from the holotype; pos-
sibly an aberrant jninctatiis in which the preocular is separated
from the upper labials by nasal and ocular).

Typhlops tettensis tettensis (Peters) Tete Blind-Snake

18(30. Onycliocephahis tettensis Peters, Monatsb. Akad. Wiss. Berlin, p.

80 : Tete, Mozambique.
Range. Kilwa and Liwale, Tanganyika Territory, southwest

to Tete, Mozambique.

Typhlops tettensis rondoensis Loveridge Rondo Blind-Snake
1942. Typhlops tettensis rondoensis Loveridge, Bull. Mus. Comp. Zool.,
91, p. 25(5: Xchingidi, Rondo Plateau, Lindi district, Tanganyika
Territory.

Range. Rondo Plateau, Tanganyika Territory.

Typhlops unitaeniatus unitaeniatus Peters
Stripe-backed Blind-Snake
1878. Typhlops (Letheoiia) unitaeniatus Peters, Monatsb. Akad. Wiss.
Berlin, p. 205, pi. ii, fig. 5: "Taita," i.e. Teita, Kenya Colony.
Range. Somalia (where the race ataeniatus Boulenger also
occurs), south through coastal Kenya Colony to Buhuri, north-
east Tanganyika Territory.

Typhlops platyrhynchus Sternfeld Tanga Blind-Snake^^

1910. Typhlops platyrhynchus Sternfeld, Mitt. Zool. Mus. Berlin, 5, p.

69 : Tanga, Tanganyika Territory.
Range. Tanganyika Territory (known only from the type

locality).

Typhlops graueri Sternfeld Lake Tanganyika Blind-Snake

1912. Typhlops graueri Sternfeld, Wiss. Ergebn. Deutsch. Zentral-
Afrika-Exped. 1907-1908, 4. p. 264: Virgin forest behind bound-
ary mountains on northwest of Lake Tanganyika, Belgian Congo.

Range. Ujiji, Tanganyika Territory, northwest to Medje,
Belgian Congo.

Typhlops gracilis Sternfeld Urungu Blind-Snake

1910. Typhlops gracilis Sternfeld, Mitt. Zool. Mus. Berlin, 5. p. 70: Kit-
ungulu, Urungu, Tanganjoka Territory,

9» Typhlops opisthopachya Werner (1917), allegedly from Tanga, has been
shown by me to be a synonym of the Australian pinguia Waite.

244 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Range. Southwestern Tanganyika Territory and Northern
Rhodesia.

Typhlops pallidus (Cope) Zanzibar Blind-Snake

1868. Letheohm pallida Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 322:

Zanzibar Island.
Range. Southern Sudan; coastal belt of Kenya Colony and

Tanganyika Territory ; Pemba and Zanzibar Islands.

Typhlops uluguruensis Barbour & Loveridge
TTluguru Blind-Snake

1928. Typhlops uluguruensis Barbour & Loveridge, Mem. Mas. Conip.

Zool., 50, p. 104: Nyange, Uluguru Mountains, Tanganyika

Territory.
Range. Tanganyika Territory (known only from the type
locality).

Typhlops braminus (Daudin)

Brahniiny or Flower-Pot Blind-Snake
1803. Eryx hrami'nus Daudin, Hist. Nat. Kept., 7, p. 279: Bengal, India.
1820. Tartrix Russellii Merrem, Vers. Syst. Amphib., p. 85: India

orientali.
1838. Onychocephalus Capensis A. Smith, Illus. Zool. S. Africa, Eept., pi.

ii, fig. 3; pi. liv, figs. 9-16: South Africa ("Interior of" may

be considered to be erroneous).
1845. Argyrophis truncatus Gray, Cat. Lizards Brit. Mus., p. 138:

Philipx)ine Islands.
1860. Ophthalmidium tenue Hallowell, Proc. Acad. Nat. Sci. Philadelphia,

p. 497 (type believed lost) : Hong Kong.

1863. Typhlops inconspieuo'us Jan, Elenco Sist. Degli Ofidi, p. 11:

Madagascar.

1864. Typhlops pammeces Giinther (new name for tenue Hallowell: pre-

occupied), Kept. British India, p. 176, pi. xvi, fig. C.
1882. Typhlops (Typhlops) euproctus Boettger, Zool. Anz., 5, p. 479:

Loucoube, Nossi Be, Madagascar.
1889. Typhlops comorensis Boulenger, Ann. Mag. Nat. Hist., (6) 4,

p. 361 : Comoro Islands.
1906. Typhlops braminus var. arenicola Aimandale, Mem. Asiatic Soc.

Bengal, 1. p. 192: Bamnad, South India.
1906. Typhlops limbricM Annandale, Mem. Asiatic Soc. Bengal, 1. p.

193 : Ramnad, South India.
1909. Typhlops braminus var. pallidus Wall (not Cope:1868), Journ.

Bombay Nat. Hist. Soc, 19, p. 609: Dibrugarh, Upper Assam.
1909. Typhlops microcephalus Werner, Jahresb. Ver. Nat. Wiirttemberg

LOVERIDQE: E. AFRICAN REPTILES AND AMPHIBIANS 245

(Stuttgart), 65. p. 60: Madagascar.
1910. Glauconia braueri Stenifeld, Mitt. Zool. Mus. Berlin, 5, p. 69:
Baganioyo, Tanganyika Territory.

1918. Typhlops capenMs Rendahl (sp. n.; not of A. Smith), Arkiv. Zool.,

11. No. 17, p. 1, figs. 1-3: "Capeland," i.e. South Africa.

1919. Typhlops fletcheri Wall, Journ. Bombay Nat. Hist. Soc, 26. p.

556: Nilgiris, India.

Range. Coastal zone of Kenya Colony ; Tanganyika Territory ;
Mozambique and South Africa. On islands of Socotra ; Zanzibar ;
Comoro; Madagascar and Mauritius.^^

Southern Europe and Asia (Arabia to China) ; islands of the
Indian Ocean; China Sea; Pacific Ocean (Mariauna; Caroline;
Marshall; Solomon; Hawaiian); West Indies; South America
(Mexico to Argentina).

Typhlops lumbriciformis (Peters) Wormlike Blind-Snake

1874. OnycJioceplialus (Lethcobin) lumbriciforviis Peters, Monatsb.

Akad. Wiss. Berlin, p. 377: "Zanzibar Coast," i.e. Tanganyika

Territory.
1904. Typhlops Meebergi Werner, Zool. Anz., 27. p. 464: Usambara,

Tanganyika Territory.
Range. Coastal Kenya Colony and Tanganyika Territory;
Zanzibar Island.'""

Family LEPTOTYPHLOPIDAE
Genus LEPTOTYPHLOPS Fitzinger

1824. Stenostoma Wagler (not Latreille:1810), in Spix, Serp. Brasil.,
p. 68. Type by monotypy: S. albifrons Wagler.

1843. Leptotyphlops Fitzinger, Syst. Rept., p. 24. Type by original
designation: Typhlops nigricans Schlegel.

1843. Bucephalus Fitzinger, Syst. Rept., p. 24. Type by original designa-
tion: Typhlops bilineatus Schlegel.

99 The world-wide distribution of this tiny snake is attributable to transport in
soil surrounding the roots of introduced plants, at other times in ballast. Its
first appearance in a country is apt to lead to redescription. The above list of
synonyms is not necessarily complete. Possibly some of the five Typhlops with
20 midbody scale-rows, described from Ceylon by Taylor (1947, Univ. Kansas
Sci. Bull., 31, pp. 287-298) may be based on individual variants. The Malagasy
species rew<cri Boettger (1882), together with its synonym lemi Boettger (1882),
was removed from the synonymy by Martens in 1922.

100 The snake from "Fwambo," I.e. Fwamba, Northern Rhodesia, assigned to
lumbriciformis by Boulenger (1896, Cat. Snakes Brit. Mus., 3. p. 590), Is actually
referable to T. gracilis Sternfeld (1910) which occurs at nearby Abercorn.

246 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1844. Catodon Dumeril & Bibron (not Linnaeus: 1761), Erpet. Gen., 6,

p. .318. Type by monotypy: Typhlops septemstriatus Schneider.
1843. Glavxonia Gray, Cat. Lizards Brit. Mus., p. 139. Type by mono
typy: Typhlops nigricans Schlegel.

1845. Epictia Gray, Cat. Lizards Brit. Mus., p. 139. Type by present

designation: Typhlops undecimstriatus Schlegel = Stenostoma
albifrons Wagler.

1833. Rena Baird & Girard, Cat. N. American Kept., p. 143. Type by
subsequent designation: E. didcis Baird & Girard.

1857. Sabrina Girard, Proc. Acad. Nat. Sci. Philadelphia, p. 181. Type
monotypy : Typhlops tessellatum Tschudi 3= Stenostoma albi-
frons Wagler.

1861. R})amphostoma Jan (not Wagler: 1830), Arch. Zool. Anat. Phys.,
1. p. 190: Type by monotypy: Stenostoma macrorhynchum Jan.

1861. Tricheilostoma Jan, Arch. Zool. Anat. Phys., 1. p. 190. Type by
present designation: Stenostoma mucrolepis Peters.

1861. Tetracheilostoma Jan, Arch. Zool. Anat. Phys., 1. p. 191. Type by
monotyijy: Typhlops bilineatus Schlegel.

1881. Singonodon Peters, Sitzb. Ges. Naturf. Freunde Berlin, p. 71.
Type by original designation: Typhlops septemstriatus Schnei-
der.

1885. StfMOstomophis Eochebrune, Faune Senegambie, Kept., p. 142.
New name for Stenostoma Wagler (preoccupied).

Leptotyphlops conjuncta conjuncta (Jan)"'^
Jan's All-black AVorm-Snake
1861. Stenostoma conjunctum Jan, Arch. Zool. Anat. Fisiol., 1. p. 189:

South Africa.
1876. Stenostoma groutii Cope, Journ. Acad. Nat. Sci. Philadelphia, (2)

8. p. 128: Umvoti Mission, Natal.
1909. Glauconia merl:cri Werner, Jahres. Ver. Nat. Wiirttemberg, 65. p.

61 : Moshi, Tanganyika Territory.
1912. Glauconia latirostris Sternfeld, Wiss. Ergebn. Deutsch. Zentral-
Afrika-Exped. 1907-1908, 4, p. 264: Northwest of Lake Tan-
ganyika Territory, Belgian Congo.
Range. Uganda (Entebbe) ; Kenya Colony and Tanganyika
Territory south to Natal. ^°"

Leptotyphlops boulengeri (Boettger)
Manda Flesh-pink Worm-Snake

101 Occasionally East African specimens have been identified as distanti (Bou-
lenger) ; nigricans (Schlegel) ; ecutifrona (Peters) or signata (Jan).

102 In Transvaal this form meets with the western race L. c. distant (Bon-
lenger) of Southern Rhodesia and Bechuanaland.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 247

1913. Glauoonia houlcngeri Boettger, in Voeltzkow, Reise in Ostafrika,

3, p. 354, pi. XXV, fig. 1 : Manda Island, Kenya Colony.
Range. Manda and Lamu Islands, Kenya Colony.

Leptotyphlops emini emini (Boulenger)^"^
Emin's All-black Worm-Snake
1890. Glauoonia emini Boulenger, Ann. Mag. Nat. Hist., (6) 6, p. 91:

Karagwe [district, Bukoba, Tanganyika Territory].
Range. (Ethiopia and Somalia); southern Sudan; Uganda;
Kenya Colony; Tanganyika Territory (and Mafia Island), west
through Belgian Ruanda-Urundi and eastern Belgian Congo to
Nyamkolo, Northern Rhodesia.

Leptotyphlops emini pembae Loveridge
Pemba White-chinned Worm-Snake
1941. Leptotyphlops emini pemhae Loveridge, Proc. Biol. Soc. Washing-
ton, 54. p. 177: Wingwi Pwana, Pemba Island.
Range. Pemlm Island (abundant).

Leptotyphlops longicauda (Peters)

Lung-tailed Flesh-pink Worm-Snake
1854. Stenostoma longicaxidum Peters, Monatsb. Akad. Wiss. Berlin, p.

621 : Tete, Mozambique.
Range. ^"'Coastal Kenya Colony; Tanganyika Territory and
Mozaml)ique, east through Nyasaland to Northern Rhodesia.

Leptotyphlops fiechteri (Seortecci) White-bellied Worm-Snake
1929. Glaiiconia fiechteri Seortecci, Atti. Soc. Ital. Sci. Nat. (Milano),

68. p. 266, figs. — : Villaggio Duca degli Abruzzi, Somalia.
Range. Somalia, south to Lake Rudolf, Kenya Colony.

Family BOIDAE
Subfamily PYTHONINAE
Genus PYTHON"' Daudin

103 Ethiopian and Somali snakes have a white-tipped tail, so maj- be eub-
speciflcally distinguishable as suggested by Parker (1949).

104 Reported, with a question mark as to identification, from Somalia by
Seortecci (1939).

105 Morelia Gray (1842) is regarded as generically distinct. Simalia Gray
I1S49), Asjjidopython A. B. Meyer (1874), and Hypaspistes D. Ogilby (1891) —
all of which have Boa amethistina Schneider (1801), or synonyms of it, as type
— are synonyms of Liasia Gray (1842), to which amethistina was transferred bj
StuU (1935).

248 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1803. Python Daudin (part), Mag. Encycl. (March, An. 8), p. 434 and

Hist. Nat. Kept., 5. p. 226. Type by subsequent designation:

Coluber molurvs Linnaeus.
1830. Constrictor Wagler (not Laurenti:1768), Syst. Nat. Amphib., p.

168. Type by subsequent designation: Boa reticulata Schneider.
1842. Heleionomus Gray, Zool. Misc., p. 42. Type by monotypy: IE.

variegatus Gray = Coluber sebae Gmelin.

1842. Hortulia Gray, Zool. Misc., p. 43. Type by monotypy: Python

natalensis A. Smith = Coluber sebae Gmelin.

1843. Asterophis Fitzinger, Syst. Eept., p. 24. Type by original designa-

tion: Python tigris Daudin = Cohiber wolurii^ Linnaeus.
1884. Aspidoboa Sauvage, Bull. Soc. Philom. Paris, (7) 8. p. 143. Type
by monotypy: Python c^irtus Schlegel.

Python sebae (Gmelin) Common African Python

1789. Coluber Sebae Gmelin, Syst. Nat., ed. 13, 1. Part 3, p. 1118: (based
on Seba, Thesaurus, 2. pi. cxcix, fig. 2) "America" (error).

1789. Coluber Speciosus Bonnaterre, Encyl. Method. Regnes Nat., Ophiol.,
p. 17: "Brazil" (error).

1801. Boa Eieroglyphica Schneider, Hist. Amphib., 2, p. 266: Cairo,
Egypt (by inference as taken from Seba).

1803. Python Eouttuyni Daudin, Hist. Nat. Rept., 5. p. 254: No locality.

1833. Python 'natalensis A. Smith, S. African Quart. Journ., 2, p. 64:
Natal, South Africa.

1842. Heleionomus variegatus. Gray, Zool. Misc., p. 43 : No locality.

1845. Python Liberiensis Hallowell, Proc. Acad. Nat. Sci. Philadelphia,
p. 249: Liberia.

1936. Python jubalis Pitman, Uganda Journ., 3, p. 211: Somalia ("ma-
terial in the museum at Mogadiscio, though I am unaware of
any published description." nomen nudum).

Range. Africa south of Senegal, the Sudan and Eritrea;
including Uganda; Kenya Colony and Tanganyika Territory.

Subfamily BOINAE
Genus ERYX Daudin

1803. Eryx Daudin (part), Mag. Encycl. (March, An. 8), p. 437, and
Hist. Nat. Rept., 7, p. 251. Type by subsequent designation:
Boa turcica Olivier.

1803. Clothonia Daudin, Hist. Nat. Rept., 7, p. 283. Type by monotypy:
Boa anguiniformis Schneider = Boa johnii Russell.

1830. Gongylophis Wagler, Nat. Syst. Amphib., p. 192. Type by mono-
typy: Boa conica Schneider.

LOVERIDQE: E. AFRICAN REPTILES AND AMPHTBIANS 249

1849. Cusoria Gray, Cat. Snakes Brit. Mus., p. 107. Type by monotypy:

C. elegans Gray.
1864. Cursoria Giinther, Rept. British India, p. 333. Emendation for

Cusoria Gray.

Eryx colubrinus loveridgei Stiill Kenya Sand-Boa

1932. Eryx thehaieus loveridgei Stull, Occ. Papers Boston Soc. Nat.
Hist., 8, p. 29, pi. ii, fig. B: "Mbunyi" i.e. Mbuyuni, Kenya
Colony.
Range. Arid areas of Kenya Colony and northern Tanganyika
Territory (at Kahe).'"'

Family COLUBRIDAE

Subfamily COLUBRINAE

Genus BOTHROPHTHALMUS Peters

18(53. Bothrophthalmus Peters, Monatsb. Akad. Wiss. Berlin, p. 287.
Type by monotypy: Elaphis (Bothrophthalmus) UneaUis Peters.

Bothrophthalmus lineatus lineatus (Peters)
Red-and-Blaek Striped Snake
1863. Elaphis (Bothrophthalmus) lineatus Peters, Monatsb. Akad. Wiss.

Berlin, p. 287: Guinea.
1863. Bothrophthalmus melanozostus Jan, Elenco Sist. Degli Ofidi, p.

62 : Gold Coast.
Range. Uganda, southwest through Belgian Congo to Angola,
northwest ("but with local races in the Cameroons) to French
Guinea.

Genus LYCODONOMORPHUS Fitzinger

1843. Lycodonomorphus Fitzinger, Syst. Eept., p. 27. Type by original

designation: Coronella rufula Schlegel = Coluher rufulus Lich-

tenstein.
1848. Lycodontomorphus Agassiz, Nomen, Zool., Index Universalis,

p. 628. Emend, for Lycodonomorphus Fitzinger.
1863. Neusterophis Giinther, Proc. Zool. Soc. Loudon, p. 16, footnote.

Type by monotypy: Natrix laevissima Gunther.
1893. Ablahophis Boulenger, Cat. Snakes Brit. Mus., 1. p. 318. Type by

106 While the "Uganda" specimen in the Museum of Comparative Zoology
possibly came from that country, as it was bought from a dealer in the days of
the "Uganda Railway," It may have come from Voi where the species is relatively
common in the vicinity of the railway station.

250 BULLETIN : MUSEUM OF COMPARATm: ZOOLOGY

uionotypy: Coluber rufulns Lichtenstein.
1893. Glypholycus Giinther, Proe. Zool. Soc. London, p. 629. Type by

nionotypy: G. bicolor Giinther.
1924. Nerophidion Werner, Sitzb. Akad. Wiss. Wien, 133, Abt. 1, p. 53..

Type by nionotypy; N. hypsirhinoides Werner = Glypholycus

bicolor Giinther.

Lycodonomorphus bicolor (Giinther)

Tanganyika White-bellied Water-Snake
1893. Glypholycus bicolor Giinther, Proc. Zool. Soc. London, p. 629,

fig. — : Lake Tanganyika.
1924. Nerophidion hypsirhinoides Werner, Sitzb. Akad. Wiss. Wien, 133,

Abt. 1, p. 53: No locality.
Range. Lake Tanganyika, Tanganyika Territory; also in lake
off Northern Rhodesia ; Belgian Congo and Ruanda-Urundi.

Lycodonomorphus rufulus whytii (Boulenger)
AVhyte 's Water-Snake
1897. GlypholycH.i wliytii Boulenger, Proc. Zool. Soc., London, pp. 800,

802, pi. xlvi, fig. 2: Fort Hill, Nyasaland.
Range. Southern Tanganyika Territory and northern Nyasa-
land, south ( ? down eastern side of Lake Nj'asa) to the Zambezi,
Mozambique.

Genus BOAEDON Dumeril & Bibron

1854.1°^ Boaedon Dumeril & Bibron, Erpet. Gen., 7, p. 357. Type by

present designation : B. lineatum Dumeril & Bibron.
1854. Eugnathus Dumeril & Bibron, Erpet. Gen., 7, p. 406. Type by

nionotypy : Lycodon geometricus Schlegel.
18.14. Alopecion Dumeril & Bibron, Erpet. Gen., 7. p. 416. Type by

nionotypy: A. annulifer Dumeril & Bibron = Lycodon guttatus

A. Smith.
1856. Holuropholis A. Dumeril, Revue Mag. Zool. (2), 8. p. 465. Type

by mouotypy: E. oUvaceus A. Dumeril.
1885. Catapherodon Roehebrune, Faune Senegambie, Rept., p. 186. Type

by present designation: Boaedon unicolor Dumeril & Bibron =

Lycodon fuligvnosus Boie.
1893. Theleu.s Cope, American Nat., 27, p. 482. Type by monotypy:

Coelopeltis virgaia HaUowell.

107 Boulenger (1S93, Cat. Snakes Brit. Mus., 1, p. 327) gives 1853 when the
Mem. Acad, ^^cl. Paris, 23, pp. 399-536 (reprint pp. 1-140), pis. i-ii, published M.
DuiiiSril's "Prodoaie de la Classification des Reptiles Ophidiens." But A. M. C.
Dumeril there (p. 460) used only Bo6dou, the French form of the word, as was
also the case with Eugnathes (p. 461) and Alop^cion (p. 462).

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 251

Boaedon fuliginosus fuliginosus (Boie)^°^
Coniiiioii House-Snake
1827. Lycodon fuliginosus Boie, Isis von Oken, 20, col. 551: "Java"

(error).
1837. Lycodon unicolor Scblegel (uot Boie), Essai Phys. Serpents, 1, p.

142; 2. p. 112: Guinea Coast.
1854. Boaedon Lincatum Dumeril & Bibron, Erpet. Gen., 7, p. 363:

Gold Coast.
1857. Boaedon quadriviUatum Hallowell, Proc. Acad. Nat. Sci. Phila-
delphia, p. 54: Isles de Lo.s, off French Guinea.
1859. Boaedon quadrilineatum A. Dumeril, Arch. Mus. Hist. Nat.

(Paris), 10. p. 193, fig. 4: Bissau, Portuguese Guinea.
1867. Alopccion variegatum Bocage, Jorn. Sci. Lisboa, 1, p. 230: Ben-

guela and Novo Redondo, Angola.
1888. Boodon bipraeo&ularis Giinther, Ann. Mag. Nat. Hist., (6) 1, p.

330, pi. xviii, fig. B: Lake Tanganyika; and Rabai Hills, Kenya

Colony.
1902. Boodon Uneatus var. plutonis Werner, Verb. Zool. Bot. Ges. Wien,

52, p. 334: Boke, Rio Nunez, French Guinea.
1932. Boaedon maculatus Parker, Proc. Zool. Soc. London, p. 363: Bihen,

1500 feet, 8° 25' N., 48° 25' E., British Somaliland.
Range. Eritrea, south through Uganda; Kenya Colony; Tan-
ganyika Territory ; Pemba ; Zanzibar and Mafia Islands to Natal,
west to the Cape, north (except for Southwest Africa where B. f.
mentalis Giinther occurs) and west to Senegal east to Eritrea.

Boaedon olivaceus (Dumeril) Gaboon Water-Snake

1856. Holuropholis olivaceus A. Dumeril, Revue Mag. Zool. (2), 8, p.

466 : Gabon, French Congo.
Range. Uganda, southwest through the Belgian Congo to

Angola, north and west to French Guinea.

Genus LYCOPHIDION Fitzinger

1843. Lycophidion Fitzinger, Syst. Rept., p. 27. Type by original

designation: Lycodon horstoTcii Schlegel = L. capensis A. Smith.

1844. Lycophydion Agassiz, Nomen. Zool., Rept., p. 27. Emendation for

los Uufortuuately there seems to be no way of avoiding the unwelcome fact
that this uaine should be used for what is probably the commonest of African
snakes. In part the species owes its lengthy synonymy to the variability of its
coloration, the young usually differing from the adults : though at times the
adults may reta'iu the juvenile coloring, they are generally bright brown, dark
brown or black. So far I have failed to separate an East African race, but
trinomials are necessary as races occur in the rain-forests of West Africa
{B. j. virgatus (Hallowell) Cameroous to Liberia), arid areas of Southwest
Africa (jB. /. mentalis Giinther), and also of Arabia (B. /. arabicua Scortecci : if
distinct).

252 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Lycophidion Fitzinger : 1843.
1857. Lycophidium Agassiz, Nomen. Zool., Index Univers., p. 218.
Emendation for Lycopliydion Agassiz: 1844.

1857. LissopMs Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p. 59.

Type by original designation: L. laterale Hallowell.

1858. Metoporhina Giinther, Cat. Snakes Brit. Mus., p. 197. Type by

monotypy: Coluber irroratus Leach.

Lycophidion meleagris Boulenger Speckled Wolf-Snake

1893. Lycophidivm meleagris Boulenger, Cat, Snakes Brit. Mus., 1,

p. 337, pi. xxi, fig. 2: Ambriz and Ambrizette, Angola.
Range. Usambara; Magrotto and Uluguru Mountains, Tan-
ganyika Territory, west through the Belgian Congo to Angola.

Lycophidion capense omatum Parker Ornate Wolf-Snake
1936. Lycophidion ornatum Parker, Novit. Zool. (Tring), 40, p. 122:

Congulu, Angola.
Range. Virgin forests of Uganda ; western Kenya Colony and

western Tanganyika Territory, west through Belgian Ruanda-

Urundi and the Belgian Congo to Angola, xilso Gold Coast

(M.C.Z. 49606, etc.).

Lycophidion capense uzungwense Loveridge
Red-snouted Wolf-Snake
1932. Lycophidion capense uzungioensis Loveridge, Bull. Mus. Comp.
Zool., 72, p. 375: Dabaga, Uzungwe Mountains, Tanganyika
Territory.
Range. Uplands of northwestern (at Kibondo) and southern
Tanganyika Territory.

Lycophidion capense capense ( Smith ) ^°®
Common Wolf-Snake
1831. Lycodon capensis A. Smith, S. African Quart. Journ., 1, p. 18:

' ' Kurrichane, ' ' i.e. Rustenberg district, Transvaal.
1837. Lycodon horstoMi Schlegel, Essai Phys. Serp., 2, p. Ill, pi. iv,

figs. 10-11: Cape of Good Hope in the uncultivated parts near

the frontiers of Kaffraria (restricted).
1888. Lycophidium Capense mut. midtimacidata Boettger, Ber. Sencken-

berg, Naturf . Ges., p. 67 : Povo Ncmlao and Povo Netonna, near

Banana, Belgian Congo.
1893. Lycophidium jaclcsoni Boulenger, Cat. Snakes Brit. Mus., 1. p.

340, pi. xxi, fig. 3: Mount Kilimanjaro, Tanganyika Territory;

Lamu, Kenya Colony.
1893. Lycophidium ahyssinicum Boulenger, Cat. Snakes Brit. Mus., 1,

p. 342, pi. xxii, fig. 1 : Southern Abyssinia.

LOVERIDQE: E. AFRICAN REPTILES AND AMPHIBIANS 253

Range. Egypt (Fayum only), east to Arabia, south through
Uganda; Kenya Colony; Tanganyika Territory; Pemba and
Zanzibar Islands ;^'''-* to Natal, west to the Cape, northwest to
French Guinea (possibly Portuguese Guinea and Senegal if L. c.
gamhiense Roehebrune is included).

Genus HORMONOTUS Hallowell

1857. Honnonotu^ Hallowell, Proc. Acad, Nat. Sci. Philadelphia, p. 56.
Type by monotypy : H. audax Hallowell = Lamprophis modestus
Dumeril & Bibron.

Honnonotus modestus (Dumeril & Bibron)

1854. Lamprophis modestus Dumeril & Bibron, Erpet. Gen., 7. p. 429:

Guinea.
1857. Eorvwnotus atidax Hallowell, Proc. Acad. Nat. Sci. Philadelphia,

p. 56: Gabon, French Congo.
1863. Heterolepis glabcr Jan, Elenco Sist. Degli Ofidi, p. 98: Boutry,

Ashanti, Gold Coast.
1888. Boodon (Alopecion) Vossii Fischer, Jahrb. Hamburg. Wiss. Anst.,

5, p. 3, pi. i, fig. 1 : Cameroon.
Range. Uganda, southwest through Belgian Congo to Angola,
northwest to French Guinea.

109 Parker (1949, Zool. Vorh. Rijksmus. Nat. Hist. Leiden, No. 6, pp. 54-56)
points out tliat it is doubtful whethier the types of acutirostre Giintlier (1868,
Ann. Mag. Nat. Hist., (4) 1, p. 427, pi. xix, fig. D), though sent by Sir John
Kirk jroin Zanzibar, originated there, for in a letter from Sir John (1867, Proc.
Zool. Soc. London, p. 952) he states he had a "valuable collection of snakes
and insects from Mozambique" (italics H.W.P.). The collection was received at
the British Museum iu 1868. Parker adds that since 1867 no wolf-snakes with
such low scale-counts as the cotypes of acutirostre have been taken on Zanzibar
Island. To which I might add that the sesing of some as given in the catalogue
(1893, p. 338) is in error. In removing L. c. acutirostre from this check list it
is as well to draw attention to the fact that in the non-montane coastal belt of
East Africa there is a population of cnpense which can be separated from the
more or less white-throated c. capense of the interior by their uniformly blackish-
brown throat and undersurfaces. The Museum of Comparative Zoology has such
specimens from Kibwezi ; foot of Mbololo ; Malindi ; Changamwe near Mombasa
(seen) ; Kilosa ; Morogoro ; Bagamoyo ; Zanzibar ; Dar es Salaam ; Mbanja near
Lindi ; Liwale ; Tunduru and Ugano.

Parker mentions that the British Museum specimen with the lowest counts
(9 :V.156:C.20) is from Kosi Bay, Zululaud. Presumably the same snake that
in 1908 Boulenger referred to semiannulis Peters, with the qualification that it
lacked the dark crossbars of the type from Tete, Mozambique. This was precisely
the case with the eight snakes (3 $ $, 5 9 $ :V.130-144 ;C. 24-31) that I person-
ally captured at Lumbo, Mozambique and referred to semiannulis in 1923. If
acutirostre came from Mozambique it is highly probable that the Lumbo snakes
represent the same thing. This suggests that acutirostre may be a synonym of
semiannulis which — except for a very questionable Angola record of 1898 —
has not been taken since Peters described it over a hundred years ago. I
failed to find any Lycophidion during the month that I spent at Tete in 1949.

254 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus MEHEXYA Csiki

1847. Eeterolepis A. Smith (not Nees:1834), Illus. Zool. S. Africa,

Kept., footnote to pi. Iv. Type by original designation: E.

capensis A. Smith.
1858. Simoccphahis Giinther (not Schodler:ix.l858), 9.X.1858, Cat.

Snakes Brit. Mus., p. 194. New name for Eeterolepis Smith

(preoccupied).
1903. Grohbenia Poche (not Holdhaus:28.vii.l903), 21.ix.l903, Zool.

Anz., 26, p. 699. Type by original designation : Eeterolepis

poensis A. Smith.

1903. Mehelya Csiki, 15.xi.l903, Eovartani Lapok, p. 198, footnote. New

name for Grohhenia Poche (preoccupied).

1904. Sichcnrockia Poche, 13.ix.l904, Zool. Anz., 28. p. 38. New name

for Grobbenia Poche (preoccupied).

Mehelya capensis savorgnani ( Mocquard )
Northwestern Cape File-Snake

1887. Efiterolepis Savorgnani Mocquard, Bull. Soc. Philom. Paris, (7)

11, p. 27, pi. ii, figs. 4-4b: Ogooue River, Fi-ench Congo.
1893. Simocephalus chanleri Stejneger, Proc. U. S. Nat. Mus., 16. p. 726:

Wange, opposite Manda Island, Kenya Colony.
1901. Simocephalus phyllophis Werner, Zool. Anz., 24. p. 301, figs. 3-4:

Cameroon.
1907. Simocephalus Butleri Boulenger, Ann. Mag. Nat. Hist., (7) 20.

p. 489: Between Wau and Chakchak, Bahr el Ghazal Province.

Sudan.
1910. Simocephalus unicolor Boulenger, Ann. Mag. Nat. Hist., (8) 5,

p. 512: Fort Hall, Kenya Colony.
1910. Mehelya (or Simocephalus) lamani Lonnberg, Arkiv. Zool., 7.

No. 8, p. 1 : Mukimbungu, lower Congo River, Belgian Congo.
1913. Mehelya {Simocephalus) so?naiierms Lonnberg & Andersson, Arkiv.

Zool., 8, No. 20, p. 2: "Kismayu, " i.e. Chisimaio, Somalia.
Range. Southern Sudan and southern Somalia, west through
Kenya Colony; Uganda; Belgian and French Congo to French
Cameroon.

Mehelya capensis capensis (Smith)
Southeastern Cape File-Snake
1847. Eeterolepis capensis A. Smith, Illus. Zool. S. Africa, Eept., pi. Iv:

Eastern districts of Cape Province, South Africa.
1874. Eeterolepis Gueinzii Peters, Mouatsb. Akad. Wiss. Berlin, p. 164,

pi. — , fig. 2: Port Natal, i.e. Durban, Natal.
Range. Tanganyika Territory/^" south through Nyasaland to
Natal, west thi-ough Bechuanalan<l to Angola.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 255

Mehelya poensis (Smith) Western-Forest File-Snake

1847. n.eterolcpis poensis A. Smith, Illus. Zool. S. Africa, Eept., footnote

to pi. Iv : Fernando Po.
1854. Heterolepis bicarinatus Dumeril & Bibrou, Erpet. Gen., 7, p. 422:

Fernando Po.
Range. Uganda southwest through Belgian Congo to Angohi,
northwest to French Guinea."^

Mehelya stenophthalmus (Mocquard) Small-eyed File-Snake

1887. Heterolepis stenophthalmus Mocquard, Bull. Soc. Philom. Paris,

(7) 11. p. 16, pi. i, figs. 1-lb: Assini, Gold Coast and Cape
Lopez, French Congo.
1911. Gonionotophis microps Boulenger, Ann. Mag. Nat. Ilist., (8) 8,

p. 370: Bitye, French Cameroon.
Range. Uganda, west through Belgian Congo to Gold Coast ;
Portuguese Guinea.

Mehelya nyassae (Giinther) Nyasaland File-Snake

1888. Simocrphalus nyassae Giinther, Ann. Mag. Nat. Hist., (6) 1, p.

328: Lake Nyasa, Nyasaland.
1906. Gonionotophis degrijisi Werner, Zool. Anz., 30, p. 53: Usambara

Mountains, Tanganyika Territory.
Range. Kenya Colony (Tana River :A.L.) ; Tanganyika Ter-
ritory and Zanzibar Island, south through Mozambique, Nyasa-
land and Southern Rhodesia to the Transvaal and Natal. Also
Belgian Ruanda-Urundi.

Genus GEODIPSAS Boulenger "^

1896. Gcodipsas Boulenger, Cat. Snakes Brit. Mus., 3. p. 32. Type by
present designation: Tachymcni^ infraJincattts Giinther.

Geodipsas vauerocegae Tornier Usambara Forest-Snake
1902. Geodipsus vauerocegae Tornier, Zool. Anz., 25. p. 703: Usambara

Mountains, Tanganyika Territory.
Range. Virgin forests of the Usambara ; Magrotto and Ulu-

guru Mountains, Tanganyika Territory.

Geodipsas procterae Loveridge Uluguru Forest-Snake

110 Voeltzkow's listing of "poensis" from Zanzibar is assumed to have been
based on misidentifled c. capensis from coastal Tanganj-ika (= Zanzibar auct.).

111 Snakes of this genus possess (1) hjpapophyses on the posterior vertebrae:
(2) grooved maxillary teeth: (:!) a forked sulcus spermaticus. In the last two
characters they differ from Natriciteres to which they bear at least a superficial
resemblance. Other species occur in the Cameroons and in Madagascar.

256 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

1922, Geodipsas procterae Loveridge, Proc. Zool, Soc. London, p. 313:

Uluguru Mountains, Tanganyika Territory.
Range. Virgin forests of the Uluguru Mountains, Tanganyika
Territory.

Genus NATRICITERES Loveridge "'-

1953. Natrieiteres Loveridge, Bull. Mus. Comp. Zool., 110, p. 248. Type
by original designation: Coronella olivacea Peters.

Natrieiteres olivacea olivacea (Peters) Olive Marsh-Snake

1854. Coronella olivacea Peters, Monatsb. Akad. Wiss. Berlin, p. 622:

Tete, Mozambique.
1860. Coronella (Mcisodon) dumerilii Giinther, Proc. Zool. Soc. London,

p. 429, fig. — : "Sierra Leone" (error, fide Boulenger, for Gold

Coast).
1863. Enicognathus punctatosirmtus Jan, Arch. Zool. Anat. Fisiol., 2,

p. 278 : Type in Hamburg Museum, but no locality.
1877. Neusterophis atratus Peters, Monatsb. Akad. Wiss. Berlin, p. 614,

pi. — , fig. 1 : Chinchoxo, Cabinda.
1886. Grayia Guirdi Dollo, Bull. Mus. Eoy. Hist. Nat. Belgique, 4. p.

158, figs. 1-2: Lake Tanganyika region, Belgian Congo.
Range. Sudan east to Somalia, south through Uganda ; Kenya
Colony; Tanganyika Territory; Zanzibar and Mafia Islands to
southern Rhodesia, northwest through Belgian Congo to the
Gold Coast.

Natrieiteres olivacea uluguruensis (Loveridge)
Montane Marsh-Snake
1935. Natrix olivacea tduguruensis Loveridge, Bull. Mus. Comp. Zool.,

79. p. 7: Nyange, Uluguru Mountains, Tanganyika Territory,
Range. Montane forests and marshes of Tanganyika Territory,
south to southern Rhodesia, west to Angola.

Natrieiteres olivacea pembana (Loveridge)"''
Pemba Island Marsh-Snake
1935. Natrix olivacea pemhana Loveridge, Bull. Mus. Comp. Zool., 79,

p. 8: Chakechake, Pemba Island.
Range. Pemba Island.

112 The eight generic names that have been misapplied to suakes of this genus,
are listed under the original (1953) citation of Natrieiteres.

113 A possibly untenable race depending on whether a majority of Pemba snalies
have 15 midbody scale-rows : occasional individuals of the mainland montane
form have 15.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 257

Genus COLUBER Linnaeus'"

]7o8. Cohikn- Lirnnous, Syst. Nat. ed. 10, 1. p. 216: Type by designation
of Fitzinger:1843: C. constrictor Linnaeus.

1818. Scoliophis Lesueur, Journ. Phys. Chim. Hist. Nat., 86. p. 297, Type
by inonotypy: S. atlanticits Lesueur = Coluber constrictor Lin-
naeus.

J 826. Eemorrhois Boie, Isis von Okeu, 19, col. 982. Type by original
designation: Coluber hippocrepis Linnaeus.

1830. Periops Wagler, Nat. Syst, Amphib., p. 189. Type by monotypy:
Coluber hippocrepis Linnaeus (inc. Natrix bdhiensis Wagler).

1843. Uicrophis Fitzinger, Syst. Rept., p. 26. Type by original designa-

tion: Coluber inridiflavus Lacepede.

1844. Eaemorrhois Agassiz, Nomen. ZooL, Rept., p. 20. Emendation for

Eemorrhois Boie.

1853. Bascanion Baird & Giraid, Cat. N. American Rept., p. 93. Type by

subsequent designation: Colub.er constrictor Linnaeus.

1854. Coryphodon Dumeril & Bibron (part; not 0\ven:1845), Erpet.

Gen., 7, p. 180. Type by subsequent designation: Coluber capi-

stratus Liclitenstein.
1860. Platyceps Blyth, Journ. Asiatic Soc. Bengal, 29. p. 114. Type by

monotypy: P. subfasciatus Blyth =z Coluber ventromaculatus

Gray.
1862. Bascanium Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 338.

Emendation for Bascanion Baird & Girard.
1865. MegaUabes Giinther, Ann. Mag. Nat. Hist., (3) 15. p. 92. Type

by monotypy: M. olivaceus Giinther = Eerpetodryas dipsas

Schlegel.
1868. Dolichophls Gistel, in Blicke, Leben Natur Menschen (Leipzig),

p. 155. Type (fide Mertens:1936) : Coluber caspius Gmelin.

11* Coluber, as emplo.yed here, is substantially the equivalent of 'Zamenis
U'ngler" as used by Bouleu^^er (1893. Oat. Snakes Brit. Mas., 1, p. 379). Un-
fortunately the type of Za)iienis Wagler was desiguuted by Fitzinger (1843,
Syst. Nat., p. 26) as Z. aesculaijii Wagler, a species that author specifically
stated is the Coluber aesc-ulapii of LacepSde, not of I.ilnnaeus. This C. acsculapii

Lac^p&de is conimouly regarded as a synonym of what is now l^nown as Elaphe
loiigissimus Laurenti, so that Zamenis Wagler is primarily a synonym of Elaphe
Fitzinger (1S33, in Wagler, Deser. Icon. Amphib., text to pi. xxvii), whose type
is parreysil Fitzinger = Coluber quatuorliiieatus Lac6p&iie.

Siibsequent designation of yutrLv gtmonensix Laurenti as a tvpe for Zatmnis
Wagler, by Ortenburger (1U128), Malcolm Smith (1943), also Inger and Clark
(1943, Copeia, p. 143) is therefore untenable, and Zamenis Wagler is a synonym
of Coluber only "in part."

Probably other names should be added to those listed here in the synonymy
of Coluber. Four of them (together with "Zamenis") are regarded as subgenera
by Inger and Clark (loc. cit.). On the other hand, Chilolepis Fitzinger, 1843.
Syst. Kept., p. 20 (of which Coluber cliff urdi Schlegel is the type by original
de^-ignation) together with its synonyms Upalerosophis Jan, 1865, in be Fillipi.
Viagg. Persia, p. 356 (whose type was designated as S. microlepis Jan in 1930
by K. P. Schmidt when reviving Hpalero-iophis) and Loxodon (also Toxodon)
Jan. 1867, are removed from the synonymy as the Chilolepis group appears to
warrant generic status.

258 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1895. Acanthomlyx Cope, Trans. Amer. Philos. Soc, 18, p. 204. Type by

original designation: Coluber v.entromaculatus Gray.
1895. Tylanthera Cope, Trans. Amer. Philos. Soc, 18, p. 205. Tj-pe by

monotypy: Coluher florulentiis Geoffroy.
192-i. Argyrogena Werner, Sitzb. Akad. Wiss. Wien, 133. Abt. 1, p. 51.

Type by monotypy: A. rostrata Werner = Coluher fasciolatus

Shaw.

Coluber keniensis Parker Lake Baringo Snake

1932. Coluber keniensis Parker, Journ. Linn. Soc. London, Zool., 38, p.

220: Lake Baringo, Kenya Colony.
Range. Kenya Colony (known only from the holotype).

Coluber citernii (Boulenger) Somalia Snake

1912. Zamenis citernii Boulenger, Ann. Mus. Civ, Stor. Nat. Geneva

(3), 5, p. 331: Dolo, Somalia.
Range. Southern Somalia and northern Kenya Colony.^^^

Coluber florulentus smithi (Boulenger)"*'
Southern P'lowered-Snake
1895. Zamenis smithi Boulenger, Proc. Zool. Soc. London, p. 536, pi. xxx,

fig. 2 : Shebeli River, Somalia.
1935. Coronella semiornata fuscorosea Loveridge, Bull. Mus. Comp. Zool.,

79. p. 8: Mount Mbololo, Teita, Kenya Colony,
Range. Ethiopia and Somalia, south to Kenya Colony.

Genus MEIZODON Fischer

1856. Meizodon Fischer, Abhand. Naturw. Ver, Hamburg, 3. p, 112.
Type by monotypy: M. regularis Fischer ■=: Calamaria coromata
Schlegel.

Meizodon semiomatus semiomatus (Peters)"'

Southern Semiornate-Snake

115 Here reported from Kenya for the first time on the basis of a 5 (M.C.Z.
51691) from Maka Murri with 21 niidbody scale-rows; 219 ventrals ; a divided
anal ; 87 subcaudals : S labials, the 4th aud 5th entering the orbit. Total length
570 (432 + I08) mm. This snalie unquestionably represents citernii, previously
known only from the holotype.

116 The 1890 record of "smithii" from Assab, Eritrea, is assumed to be based
on a C. /. florulentus CJeoffroy. Conversely, the "florulentus" recorded by Lonn-
berg (1911) from just north of the Uaso Xyiro (as Guaso Nyiri), and also a
Lake Kudcdf snake in the Nairobi Museum are almost certainly C. /. smithi,
which I still think is only a race.

117 Trinomials are employed on account of a northern race — M. s. plumbeiceps
(Boettger :189o) with somalica (Scortecci :1932) and loveridgei Bogert :1940 as
synonyms — ranging from Sudan, Ethiopia and British Soraaliland south through
Somalia. Where the races actually meet remains to be defined, probably arbitrar-
ily, as the sole character separating them appears to be the predominance of 1
+ 2 temporals in the northern race, 2 + 2 in the southern. The Uganda record
of Boulenger (1902) is extremely questionable as no specimen is known.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 259

1854. Coronella semiornata Peters, Monatsb. Akad. Wiss. Berlin, p. 622:
Tete, Mozambique.

1879. Zamenis fischeri Peters, Monatsb. Akad. "Wiss. Berlin, p. 777:
Malindi, Kenya Colony.

1884. Coronella inornata Fischer, Jahrb. Hamburg. Wiss. Anst., 1,
p. 6, pi. i, fig. 2: Masailand, Kenya Colony.

1908. Coronella scheffleri Sternfeld, Sitzb. Ges, Naturf. Freunde Berlin,
p. 93 : Kibwezi, Kenya Colony.

1935. Coronella semiornata var. viossamhica Cott, Proc. Zool. Soc. Lon-
don, 1934, p. 967: Cbarre and Fambani, Mozambique.

Range. Uganda ; Kenya Colony and Tanganyika Territory,
south through Mozambique to Nyasaland and the Rhodesias.

Meizodon coronatus (Schlegel) Western Crowned-Snake

1837. Calamariu coronata Schlegel, Essai Phys. Serp., 2, p. 46: Gold

Coast.
18.36. Meizodon regulari^ Fischer, Abhand. Nat. Ver, Hamburg, 3, p.

112, pi. iii, fig. 3: Peki, Gold Coast.
1858. Coryplwdon margaritiferus Giinther, Cat. Snakes Brit. Mus., p.

109 : West Africa.
1860. Coronella (Meizodon) iitorqnata Giinther, Proc. Zool. Soc. Londou,

p. 428, fig. — : Senegal.
1863. Coronella elegans .Jan, Arch. Zool. Anat. Fisiol., 2, p. 255: No

locality.
1917. Zamenis tchadensis Chabanaud, Bull. Mus. Hist. Nat. (Paris), 22,

p. 451, figs. 3-5: Koalem, French West Africa.
1933. Coronella r.egularis praeornata Angel, Serpentes Afrique Occ.

Fran^aise, p. 123, figs. 45a-b: French Sudan and Uganda.
1951. Meizodon coronata camerunensis Monard, Mem. Inst. Franc.

Afrique Noire (Sci. Nat.), No. 1, pp. 151, 161: Konn, French

Cameroon.
Range. Kenya Colony and Uganda, west in equatorial belt
(5° S. to 15° N.) to Senegal.

Genus PHILOTHAMNUS Smith "«

1847. Philothamnns A. Smith, Illus. Zool. S. Africa, footnote to text for
pi. lix. Type by monotypy: Dendrophis (Philothamniis) semi-
variegata A. Smith.

1857. Chlorophis Hallowell, Proc. Acad. Nat, Sci. Philadelphia, p. 53.
Type by monotypy: C. heterodermus Hallowell.

1866. Eerpetaethiops Giinther, Ann. Mag. Nat. Hist., (3) 18. p. 27. Type

118 Chlorophis Hallowell is regarded as a subgenus, usually recognizable by the
absence of a sharply angular subcaudal keel along either side of the tail.

260 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

by monotypy: H. bellii Giinther = Chlorophis heterodermus

Hallowell.
1868. Chlorophis Theobald (not Hallowell), Cat. Rept. Asiatic Soc. Mus.,

p. 49. Type by monotypy: C. oldhami Theobald = Ahaetvlla

hoplogaster Giinther.
1895. Oligolepis Boulenger, Ann. Mag. Nat. Hist., (6) 16, p. 171. Type

by monotypy: 0. macrops Boulenger.

Philothamnus macrops (Boulenger) Usambara Green-Snake
1895. Oligolepis macrops Boulenger, Ann. Mag. Nat. Hist., (6) 16, p.

171: Usambara Mountains, Tanganyika Territory.
Range. Coastal Tanganyika Territory and Zanzibar Island.

Philothamnus heterodermiis carinahis (Andersson)
Thirteen-scaled Green-Snake
1901. Chlorophis carinatus Andersson, Svenska Vetensk.-Akad. Hand.,

27, No. 5, p. 9: Mapanja, British Cameroon.
Range. Western Kenya Colony and Uganda, west through
Belgian Congo to Fernando Po.

'■o'

Philothamnus hoplogaster (Giinther)
Southeastern Green-Snake
1863. Ahnetulla hoplogaster Giinther, Ann. Mag. Nat. Hist., (3) 11, pp.

284, 286: "Port Natal," i.e. Durban, Natal.
1866. Philothamnus n.eglectus Peters, Monatsb. Akad. Wiss. Berlin, p.

890, footnote: "Prazo Boror, " i.e. Boror, Mozambique.
1868. Chlorophis Oldhami Theobald, Cat. Rept. Asiatic Soc. Mus., p. 49:

" Simla, India " (error).
Range. Tanganyika Territory (south of the Usambara Moun-
tains), south through Mozambique and the Rhodesias to Port
Elizabeth, Cape Province, South Africa.

Philothamnus irregiilaris battersbyi Loveridge"^
Northeastern Green- Snake
1951. Philothamnus irregidaris battersbyi Loveridge, Bull. Mus, Comp.
Zool., 106. p. 51: Sipi Forest at 6000 feet on Elgon Mountain,
eastern Uganda.
Range. Sudan (east of the Nile), east through Ethiopia to
Somalia, south through Uganda and Kenya Colony to (and in-
cluding) the Usambara Mountains, Tanganyika Territory, west
to Lake Victoria.

119 This is the common green-snake of the northeastern savannas which, with
the preceding has been so long erroneously known as Chlorophis neglecttis
(Peters).

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 261

Philothomnus irregularis irregidaris (Leach)
Western Green-Suake
1789. Coluber Caeruleus LacepSde (not Linnaeus), Hist. Nat. Serpens,

2. Table Methodique, p. 100, text p. 276: Cap Vert, Senegal.
1802. Coluber caei-ulesccns Daudin (not Linnaeus: new name for caeru-
leus Laccpede), Hist. Nat. Eept., 7, p. 54.

1818. Coluber caesius Cloquet^^" (new name for caeruleus Lacepede),

Diet. Sci. Nat. (Paris) 11. p. 201.

1819. Coluber irregularis Leacli, in Bowdich, Miss. Ashantee, App. p.

494: "Fantee," i.e. Fanti, Ashanti, Gold Coast,
1843. Dendrophis Chenonii Reinhardt, Dansk. Vidensk. Selsk. Skrift., 10,

p. 246, pi. i, figs. 13-14: Guinea.
1848. Dendrophis (Philothamnus) albo-variata A. Smith, lUus. Zool.
S. Africa, Rept., pi. Ixiv, figs. 3-3b; pi. Ixv: South Africa
towards Tropic of Capricorn (also Gold Coast; Sierra Leone;
Gambia).
1875. Fhilothamnus irregularis var. longifrenatus Buchholz and Peters,

Monatsb. Akad. Wiss. Berlin, p. 199: Cameroon.
1882. Philothavinus angolensis Bocage, Jorn. Sci. Lisboa, 9. p. 7: Capan-

gombe, Angola.
1888. AJiaetulla emini Giinther, Ann. Mag. Nat. Hist., (6) 1, p, 51:

' ' Monbuttu, ' ' i.e. Mangbetu, Belgian Congo.
1888. Ahaetulla shirana Giinther, Ann. Mag. Nat. Hist., (6) 1, p. 326:

Blantyre Mission, Shire River, Nyasaland.
1893. Philothamnits Giintheri Pfeffer, Jahrb. Hamburg. Wiss. Anst., 10.

p. 85, pi. i, figs. 3-5: Quilimane, Mozambique.
1932. Chlorophis vernayi FitzSimons, Ann. Transvaal Mus., 15. p. 38:

Maun, Thamalakane River, British Bechuanaland.
Range. Tn addition to western Uganda and western Tan-
ganyika Territory: West and Central Africa south of 15° N.
and north of 20° S. but following the Zambezi to the East Coast.
More specifically: Senegal east to Metemma (introduced) on
the White Nile south down the Rift Valley (Lakes Victoria,
Tanganyika and both sides of Nyasa) into Southern Rhodesia,
west to Damaraland, northwest to Senegal.

Philothomnus heterolepidotus (Giinther) Slender Green-Snake
1863. Ahaetulla heterolepidoia Giinther, Ann. Mag. Nat. Hist., (3) 11,

p. 286 : Africa.
1888. Ahaetulla gracillima Giinther, Ann. Mag. Nat. Hist., (6) 1, p. 326:

Lower Congo River, Belgian Congo.

120 Suppressed by the International Commission on Zoological Nomenclature In
1955. See Opinions and Declarations, 9, pp. 299-308.

262 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1910. ChloropJii^ gracilis Sternfeld, Mitt. Zool. Mus. Berlin, 5, p. 64:

Dodo, French Cameroon.
1912. Chlorophis schubotsi Sternfeld, Wiss. Ergebn. Deutsch. Zentral-

Afrika-Exped. 1907-1908, 4, p. 269, fig. 4: Bwanja, near Bukoba,

Tanganyika Territory.
1923. Chlorophis bequaerti Schmidt (part), Bull American Mus. Nat.

Hist., 49, p. 7.1, fig. 3: Niangara, Belgian Congo (based on an

aberrant heterolcpidotus $ -with entire anal ^ the holotype;

omit paratype $ , a similarly aberrant irregularis) .
Range. Southern Sudan ; Uganda and western Kenya Colony,
southeast to mouth of the Rovuma River, Tanganyika Territory ;
( ? Mozambique) ; west to Angola and northwest to Togo.

Philothamnus semivariegatus dorsalis (Bocage)^^^
Stri])ed Wood-Snake
1866. Leptophis dorsalis Bocage, Jorn. Sci. Lisboa, 1. pp. 48, 69:

Molembo, Loanda Coast, Angola (restricted by Bocage: 1882).
Range. Angola, north to the Belgian and French Congo.

Philothamnus semivariegatus semivariegatus (Smith)
Spotted Wood-Snake
1847. Bendrophis (Philothamnus) semivariegata A. Smith, Illus. Zool.
S. Africa, pis. lix; Ix; Ixiv, figs, la-lb: Bushman Flat, Cape
Province, South Africa (restricted by Bogert:1940).
1866. Philothamnus punctatus Peters, Monatsb. Akad. Wiss. Berlin, p.
889: Zanzibar Coast, i.e. Tanganyika Territory.

1868. Ahaetidla Kirkii Giinther, Ann. Mag. Nat. Hist., (4) 1. p. 424:

Zanzibar Island.

1869. Bendrophis melanostigma Jan, Icon. Gen. Ophid., livr. 32, pi. ii,

fig. 3 : Mozambique.

1882. Philothamnus Smithii Bocage, Jorn. Sci. Lisboa, 9, p. 12, fig. 5:
Bissau, Portuguese Guinea (hereby restricted A.L.).

1888. Ahaetulla Bocagii Giinther, Ann. Mag. Nat. Hist., (6) 1. p. 326:
Angola.

1893. Philothamnus punctatus var. sansibaricus Pfeffer, Jahrb. Ham-
burg. Wiss. Anst., 10, p. 83 : Zanzibar Island.

Range. Includes Uganda; Kenya Colony; Tanganyika Terri-
tory; Pemba, Zanzibar and Mafia Islands. Africa south of 16° N.
except forested areas bordering the Gulf of Guinea, certain

121 It Is very doubtful if this form should be admitted to the East African
list. It is included here to invite attention to the population (of which I took
22 specimens) around Mwaya, Lake Nyasa, Tanganyika Territory. In this series^
the temporal arrangement was predominantly that of dorsalis, though in markings
they resemble typical semivariegatus in lacking the vertebral stripe which is ap-
parently not constant even among West African dorsalis.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 263

islands therein and a vague coastal area from French Congo to
Angola occupied by races, besides being absent from the Cape
Peninsula region. More specifically Gambia east to Eritrea
(though not recorded from Ethiopia), south to Natal and adja-
cent Cape Province, west and northwest (except for regions
occupied by races) to Gambia.

Genus HAPSIDOPHRYS Fischer

1856. Hapsidophrys Fischer (part), Abhand. Nat. Ver. Hamburg, 3. p.
110. Type by present designation: H. lineatus Fischer.

Hapsidophrys lineata Fischer Black-lined Green-Snake

1856. Hapsidophrys lineatus Fischer, Abhand. Nat. Ver. Hamburg, 3.

p. Ill, pi. ii, fig. 5: Elmina, Gold Coast.
1909. Gastropyxis orientalis Werner, Jahresb. Ver. Nat. Wiirttemberg,

(Stuttgart), 65, p. 56: Tanganyika Territory.
Range. Western Kenya Colony and Tanganyika Territory,
west through Uganda, southwest to Angola, north and west to
French Guinea.

Genus GASTROPYXIS Cope

1861. Gastropyxis Cope, Proe. Acad. Nat. Sci. Philadelphia, 1860, pp.

556, 558. Type by monotypy: Dendrophis smaragdina Schlegel.

Gastropyxis smaragdina (Schlegel) Keel-scaled Green-Snake
1837. Dendrophis smaragdina Schlegel, Essai Phys. Serp., 2. p. 237:

Gold Coast.
1844. Leptophis gracilis Hallowell, Proc. Acad. Nat. Sci. Philadelphia,
p. 60 : Liberia.

1856. Hapsidophrys caeruleus Fischer, Abhand. Nat. Ver. Hamburg, 3,

p. Ill, pi. ii, fig. 6: Elmina, Gold Coast.
Range. Uganda, southwest through the Belgian Congo to
Angola, north and west to Portuguese Guinea.

Genus THRASOPS HallowelU"

1857. Thrasops Hallowell, Proe. Acad. Nat. Sci. Philadelphia, p. 67.

Type by monotypy: Dendrophis flavigularis Hallowell.

1862. Ehamnophis Giinther, Ann. Mag. Nat. Hist, (3) 9. p. 129. Type

122 Rhamuophis is at best but a subgenus of Thrasops, the differences between
the two being slight, as may be seen by comparing their respective characters
given in my revision (1944, Bull. Mus. Comp. Zool., 95, pp. 124-138).

264 buTjLEtin : museum of comparative zoology

by monotypy: R. aethiopissa Giinther.
1864. Crypsidomus Giinther, Proc. Zool. Soc. London, p. 309. Substitute
name for Bhamnophis Giinther.

Thrasops jacksonii jacksonii Giinther
Western Black Tree-Snake
1895. Thrasops Jadcsonii Giinther, Ann. Mag. Nat. Hist., (6) 15, p.

528 : Kavirondo, Kenya Colony.
1905. Thrasops Bothschildi Mocquard, Bull. Mus. Hist. Nat. (Paris),

11, p. 287: "Afrique orientale anglaise, " i.e. Kenya Colony.
Range. "Western Kenya Colony and Tanganyika Territory
(west of Lake Victoria), west through Uganda to the Belgian
Congo.

*o^

Thrasops jacksonii schmidti Loveridge
Eastern Black Tree-Snake
1936. Thrasops jaclcsonii schmidti Loveridge, Proc. Biol. Soc. Washing-
ton, 49, p. 63 : Meru Forest, Mount Kenya, Kenya Colony.
Range. Central Kenya Colony (virgin forests of Mount Kenya
and Muthaiga near Nairobi).

Thrasops aethiopissa elgonensis (Loveridge)

1929. Ehamnophis aethiopissa elgonensis Loveridge, Bull. TJ. S. Nat.

Mus., 151, p. 24: Yala (= Lukosa) Eiver, Kenya Colony.
Range. Western Kenya Colony to western Uganda.

Genus SCAPHIOPHIS Peters

1870. Smphiophis Peters, Monatsb. Akad. Wiss. Berlin, p. 644. Type
by monotypy: S. aliopunctatus Peters.

Scaphiophis albopunctatus albopunctatus Peters
Gray Beaked-Snake
1870. Scaphiophis albopunctatus Peters, Monatsb. Akad. Wiss. Berlin,

p. 645, pi. i, fig. 4: Kita, French West Africa.
Range. Uganda; Kenya Colony and Tanganyika Territory,
west through Belgian Congo to the French Soudan.

Genus PROSYMNA Gray

1849. Prosymna Gray, Cat. Snakes Brit. Mus., p. 80. Type by monotypy:

Calamaria meleagris Reinhardt.
1849. Temnorhynchus A. Smith (not of Hope: 1837), Illus. Zool. S.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 265

Africa, Eept., App., p. 17. Type by monotypy: T. sundewallii
(misprint for sundevallii) A. Smith.
1863. Ligonirostra Cope, Amer. Journ. Sci. Arts, (2) 35. p. 457. New
name for Temnorhynchus Smith (preoccupied in coleoptera).

1896. Asthenophis Boulenger, Ann. Mus. Civ. Stor. Nat. Geneva, (2) 17,

p. 12. Type by monotypy: A. ruspolii Boulenger.

1902. Pseudoprosymna Lindholm, in Lampe, Jahrb. Nassau Ver. Naturk.
(Wiesbaden), 55. p. 57. Type by monotypy: P. 'bergeri Lind-
holm.

1909. Stenorhahdium Werner, Jahresb. Ver. Nat. Wiirttemberg, 65, p. 59.
Type (believed lost) by monotypy: S. temporale Werner.

Prosymna pitmani Battersby Multi-scaled Shovel-snout

1951. Prosyvina pitmani Battersby, Ann. Mag. Nat. Hist, (12) 4. p.

828: Kilwa, Southern Province, Tanganyika Territory.
Range. Southeastern Tanganyika Territory.

Prosymna ambigua bocagii Boulenger
Central African Shovel-snout

1897. Prosymna Bocagii Boulenger, Ann. Mag. Nat. Hist., (6) 19, p.

278, figs. — : Zongo, Ubangi Eapids, Belgian Congo.
Range. Southern Sudan and Uganda to northern and eastern
Belgian Congo.

Prosymna ambigua stuhlmanni (Pfeffer)
East African Shovel-snout
1893. Ligonirostra Stuhlmanni Pfeffer, Jahrb. Hamburg. Wiss. Anst.,

10. p. 78, pi. i, figs. 8-10: Usambara, Tanganyika Territory.
1906. Prosymna Vassei Mocquard, Bull. Mus. Hist. Nat. (Paris), 12. p.

250 : Mozambique.
1909. Prosymna variahilis Werner, Jahresb. Ver. Nat. Wiirttemberg, 65.

p. 57: Moshi, Tanganyika Territory.
1909. Stenorhabdium temporale Werner, Jahresb, Ver. Nat. Wiirttem-
berg, 65. p. 60: "East Africa."
Range. Coastal Kenya Colony; Tanganyika Territory and
Zanzibar Island, south through Mozambique, Nyasaland and the
Rhodesias to Zululand, Natal.

Prosymna ambigua ornatissima Barbour & Loveridge
Ornate Shovel-snout
1928. Prosymna ornatissima Barbour & Loveridge, Mem. Mus. Comp.
Zool., 50. p. 120, col. pi. ii, fig. 2: Nyange, Uluguru Mountains,
Tanganyika Territory.
Range. Uluguru Mountains Tanganyika Territory.

266 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus PSEUDASPIS Fitzinger

1843. Pseudaspis Fitzinger, Syst. Kept., p. 25. Type by original designa-
tion: Coluber canus Linnaeus.

1868. Cadmus Theobald (not Bory:1824), Cat. Rept. Asiatic Soe. Mus.,
p. 58. Type by monotypy: C. cuneiformis Theobald = Cohiber
eanus Linnaeus.

1882. Ophirhina Bocage, Jorn. Sci. Lisboa, 8, p. 300. Type by monotypj"^:
0. anchietae Bocage = Cohiher canus Linnaeus.

Pseudaspis cana (Linnaeus) Mole Snake

1758. Coluber canus Linnaeus, Syst. Nat., ed. 10, 1. p. 221: "in Indiis, "

i.e. Africa.
1768. Cohiber elegantissimus Laurenti, Syn. Eept., p. 96: (based on

Seba, Thesaurus, 1, pi. Ixxxi, fig. 9) No locality.
1789. Coluber ocellatus Gmelin, Syst. Nat., ed. 13, 1. Part 3, p. 1113:

(based on Seba, Thesaurus, 2. pi. i, figs. 3 and 8) "Zeylon and

Sina" (error).
1868. Cadmus cuneiformis Theobald, Cat. Eept. Asiatic Soe. Mus., p. 58:

"Simla, India" (error).
1872. Coronella phocarum Giinther, Proc. Zool. Soe. London, p. 836:

Robben Island, Cape Province, South Africa.
1882. Ophirhina Anchietae Bocage, Jorn. Sci. Lisboa, 8, p. 300: Caconda.

Benguela, Angola.
Range. Uplands of Kenya Colony and Tanganyika Territory,
south to Natal, west to the Cape, north through Southwest Africa
to Angola east through southern Belgian Congo.

Genus DUBERRIA Fitzinger

1826. Duberria Fitzinger (part), Neue Class. Eept., pp. 29, 55. Type
by tautonomy: Coluber arctiventris Daudin (1803) = Coluber
Duberria Merrem (1790) = Coluber lutrix Linnaeus (1758).

1830. Homalosoma Wagler, Nat. Syst. Amphib., p. 190, footnote. Substi-
tute name for Duberria Fitzinger.

1833. Braehyblemma Wagler, Isis von Oken, col. 904. Type by mono-
typy: Coluber duberria Merrem = Coluber lutrix Linnaeus.

Duberria lutrix abyssinica (Boulenger) Abyssinia Slug-eater
1894. Homalosoma abyssinicum Boulenger, Cat. Snakes Brit. Mus., 2,

p. 27G, p]. xiii, fig. 2, Lake Ashanti, Ethiopia.
1912. Homalosoma lutrix var. atrireniris Sternfeld, Wiss. Ergebn. Deut.

Zentral-Afrika-Exped. 1907-1908, 4, p. 271: "Kissenje," i.e.

Kisenyi, Belgian Ruanda.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 267

Range. Highlands of Ethiopia; Uganda; Kenya Colony and
northern Tanganyika Territory/-^ west through Belgian Ruanda-
Urundi to the Belgian Congo.

Duberria lutrix shirana (Boulenger) Shire Slug-eater

1894. Homalosoma sliiranum Boulenger, Cat. Snakes Brit. Mus., 2. p. 276,

pi. xiii, fig. 1: Shire Highlands, Nyasaland.
Range. Highlands of southern Tanganyika Territory and

Nyasaland.

Genus GRAYIA Giinther

1857. Eeteronotus Hallowell (not Laporte:1832), Proc. Acad. Nat. Sci.

Philadelphia, p. 67. Type by monotypy: Coronella triangularis
Hallowell = Coluber smyihii Leach.

1858. Gratia Giinther, Cat. Snakes Brit. Mus., p. 50. Type by monotypy :

G. silurophaga Giinther = Coluber smythii Leach.

1862. Glaniolestes Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 191.

New name for Eeteronotus Hallowell (preoccupied in Hemi

ptera).
186.'5. Leionotus Jan (not of Cocteau and Bibron:1843), Elenco Sist.

Degli Ofidi, pp. 68, 74. Type by monotypy: L. Schlegeli Jan

(based on ms. label Tropidonotus glaber in Leyden Museum).
1866. Macrophis Bocage, Jorn. Sci. Lisboa, 1, p. 67. Type by monotypy:

M. ornatus Bocage.

1863. Xenurophis Giinther, Ann. Mag. Nat. Hist., (3) 12. p. 357. Type

by monotypy : X. caesar Giinther.

Grayia smythii (Leach) Smyth's Water-Snake

1818. Coluber Smyihii Leach, in Tuckey, Explor. River Zaire, App., p.
409 : ' ' Embomma, ' ' i.e. Boma, Belgian Congo.

1854. Coronella triangularis Hallowell, Proc. Acad. Nat. Sci. Philadel-
phia, p. 100: Liberia.

1858. Grayia silurophaga Giinther, Cat. Snakes Brit. Mus., p. 51: West
Africa.

1863. Lejonotus Schlegeli Jan, Elenco Sist. Degli Ofidi, p. 74: Ashanti.
Gold Coast.

Range. Rivers and lakes of Sudan and Uganda, southwest to
Angola, north and west to Senegal.

Grayia tholloni Mocquard Tholloni's Water-Snake

1897. Grayia Tholloni Mocquard, Bull. Soe. Philom. Paris, (8) 9. p. 11:

French Congo.

1-3 Though "Zanzibar" is included on a recent list, I cannot believe that this
upland form (whicli occurs on Kilimanjaro and the Usambara Mountains) ever
L'anie from Zanzibar Island.

268 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1901. Grayia fasciata Boulenger, Ann. Mus. Congo, (1) 2, p. 9, pi. iii,
fig. 3 : Southwest of Lake Tanganyika, Belgian Congo.

Range. Rivers and lakes of Sudan; Uganda; western Kenya
Colony and western Tanganyika Territory ; west through the
Belgian Congo to Angola.

Genus BOIGA Fitzinger

124

O^

1825. Ibiba Gray, Ann. Pliilos. (2), 10. p. 209. Tjipe by monotypy:

Coluber irregularis Merrem.

1826. Boiga Fitzinger (part), Neue Class. Eept., pp. 29, 60. Type by

subsequent designation : Coluber irregularis Merrem.

1843. Bipsadomorphus Fitzinger, Syst. Eept., p. 27. Type by original
designation: Coluber trigonatus Schneider.

1843. Cephalophis Fitzinger, Syst. Kept., p. 27. Type by original desig-
nation : Dipsas dendrophila Boie.

1843. Eudipsas Fitzinger, Syst. Eept., p. 27. Type by original designa-
tion: Dipsas cynodon Boie.

1843. Macrocephalus Fitzinger, Syst. Eept., p. 27. Type by original

designation : Dipsas draincsii Boie.
1854. Opetiodon Dumeril & Bibron, Erpet. Gen., 7. p. 90.5. Type by

monotypy: Dipsas cynodon Boie.
1854. Triglyphodon Dumeril & Bibron, Erpet. Gen., 7. p. 1069. Type by

present designation : Coluber irregularis Merrem.
18.'i7. Toxicodryas Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p. 60.

Type by monotypy: Dipsas blarulingii Hallowell.
1877. Pappophis Macleay, Proc. Linn. Soc. N. S. Wales, 2, p. 39. Type

by monotypy: P. laticeps Macleay =: Coluber irregularis Merrem.
1894. Liophallus Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 427. Type

by original designation: Dipsas fuscus Gray.

Boiga blandingii (Hallowell) Blanding's Tree-Snake

1844. Dipsas Blandingii Hallowell, Proc. Acad. Nat. Sci. Philadelphia,

p. 170: Liberia.
1854. Triglyphodon fuscus Dumeril & Bibron, Erpet. G6n., 7, p. 1101 :

Ivory Coast.
1856. Dipsas fasciata Fischer, Abhand. Nat. Ver. Hamburg, 3. p. 84,

pi. iii, fig. 5 : Peki, Gold Coast.
1856. Dipsas valida Fischer, Abhand. Nat. Yer. Hamburg, 3, p. 87, pi.

iii, fig. 4: Edina, Grand Bassa County, Liberia.

124 Together with Oeodipsas (here placed near Natriciteres), the snakes from
this point to the end of the COLUBRIDAE (except for an occasional Aparallac-
tus) are backfanged. They constitute the OPISTHOGLYPHA of Boulenger, by
whom they were placed in a subfamily known as the DIPSADOMORPHINAE
(BOIGIXAB of later authors), a group since discarded as unnatural because of
independent derivation.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 269

1856. Dipsas globiceps Fischer, Abhand. Nat. Ver. Hamburg, 3. p. 89,

pi. iii, fig. 6: Edina, Grand Bassa County, Liberia.
1859. Dipsas Fischeri Jan (new name suggested by Jan to combine the

three preceding age and color variations), in A. Dumeril, Arch.

Mus. Hist. Nat. (Paris), 10. p. 212.
1871. Dipsas regalis "Schlegel" Jan, Icon. Gen. Ophid., livr. 38, note to

pi. vi, fig. 2 : Ivory Coast.
1885. Dipsas globiceps var. tumboensis F. Miiller, Verh. Naturf. Ges.

Basel, 7. p. 688: Tumbo Island, French Guinea.
Range. Uganda, southwest through the Belgian Congo to
Angola, northwest to French Guinea.^-''

Boiga pulverulenta (Fischer) Powdered Tree-Snake

1856. Dipsas pulverulenta Fischer, Abhand. Nat. Ver. Hamburg, 3. p.

81, pi. iii, fig. 1: Edina, Grand Bassa County, Liberia,
1917. Dipsadomorphu^ Boueti Chabanaud, Bull. Mus. Hist. Nat. (Paris),

1916, 22. p. 373, figs. 13-15: Porto Novo, Dahomey.
Range. Uganda, southwest through the Belgian Congo to
Angola, northwest to French Guinea.

Genus DIPSADOBOA Gtinther^^

1858. Dipsadoboa Giinther (part). Cat. Snakes Brit. Mus., p. 182.

Type by subsequent restriction: D. unicolor Giinther.
1869. Anoplodipsas Peters, Monatsb. Akad. Wiss. Berlin, p. 442. Type

by monotypy: A. viridis Peters = Dipsadoboa unicolor Giinther.

Dipsadoboa unicolor Giinther Giinther 's Green Tree-Snake
1858. Dipsadoboa unicolor Giinther, Cat. Snakes Brit. Mus., p. 183:

West Africa.
1869. Anoplodipsas viridis Peters, Monatsb. Akad. Wiss. Berlin, p. 442,

pi. — , fig. 4: "New Caledonia" (error).
1871. Eeteruru3 bicolor Jan, Icon. Gen. Ophid., livr. 38, pi. ii, fig. 3 :

No locality.
1893. Dipsadoboa assimilis Matschie, Sitzb. Ges. Naturf. Freunde Berlin,

p. 173 : Bismarckburg, i.e., Adele, Togo.
Range. Uganda, west through Belgian Congo to French

125 Reported from Zanzibar in error by Boulenger (1896, Cat. Snakes Brit.
Mus., 3, p. 77) and consequently followed by many authors.

126 For a key to the five species recognized by Laurent as of 1951, see Laurent,
1951, Revue Zool. Dot. Afr., 44, p. 211. As of 31.xii.l956 I have not seen the
description of the hii,'h-altitude Dipsadoboa unicolor viridiventris Laurent (men-
tioned by Curry-Lindahl, 1956. Ann. Mus. Royal Congo Beige, 42. p. 45, as about
to be published).

270 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Guinea (but as yet unrecorded from Nigeria: Dahomey and
Liberia)/"'

Genus TELESCOPUS Wagler

1830. Telescopus Wagler (not Telescopium Montfort:1810), Nat. Syst.

Amphib., p. 182. Type by monotyiDy: Coluber on pi. v, figs. 11-13
iu Savigny's Suppl. to Geoffioy, 1812, Descr. Egypte.

1831. Tarhophis F. L. Fleisclnnann, Dalm. Nov. Serp. Gen., p. 17. Type

by monotypy ( ? as not seen): T. fallax Fleiselimann.

1831. TrigonopMs Eiehwald, Zool. Spec. Rossiae Poloninae, Part 3, p.
175. Type by monotypy: T. iberiis Eiehwald.

1837. Ailurophis Bonaparte, Icon. Fauna Italiea, 2. Anfibi, p. 101. Type
by monotypy: Coluber vivax Schreiber = Tarbophis vivax
Dumeril & Bibron.

1846. Aelurophis Agassiz, Nomen. Zool., Index Univers., p. 9. Emenda-
tion for Ailurophis Fitzinger = Bonaparte.

193.1. MigiurtinopMs Scortecei, Ann. Mus. Civ. Stor. Nat. Geneva, 59,
p. 1. Type by monotypy: M. pulcher Scortecei.

Telescopus dhara somalicus ( Parker) ^^^
Southern Large-eyed Snake
1949. Tarbophis dhara somalicus Parker, Zool. Verb. Rijksmus. Nat.
Hist. Leiden, No. 6, p. 88: Ngatana, Tana River, Kenya Colony.
Range. Southern Somalia and northern Kenya Colony.

Telescopus semiannulatus semiannulatus Smith
Eastern Tiger-Snake
1849. Telescopus semiannulatus A. Smith, Illus. Zool. S. Africa, Rept., pi.

ixxii; South Africa (by inference).

1898. Tarbophis semiannulatus var. meridionalis Werner, Jahresb.

Abhand. Naturw. Ver. Magdeburg, 1896-1897, p. 144: South

Africa.

Range. Kenya Colony and Tanganyika Territory, south to

Natal, northwest through Basutoland and Bechuanaland to

12" Only represented in the Museum of Comparative Zoology by examples from
Uganda, Belgian Congo and the Prencli Cameroon ; nor are there any sl)ecimenl^
from west of the Cameroons in the Britisli Museum as of today.

128 The type from Xgatana was originally assigned by Boulenger (1896) to
Tarhojjhis yucntluri Anderson, considered by Parker to be synonymous with the
typical form described as Coluhcr dhara B\irskal (1775. Descr. Anim., p. 14).
Probably the two Kenya records of Tarbo/jhis ohtusus Boettger (which is a
synonym of (I. dhara) from Njiri (V Nyeri or Uaso Nyiro) by Mocquard (1902).
and Irom Lodwar by Parker (1936) are referable to T. d. somalicus. My own
early (1916-1917) records from Gobwen (now Gobuin) and Kismayu (now
Chisimaio) were made prior to the transfer of those places to what is now
^onuilia.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 271

Southwest Africa (Avhere it meets with a subspecies), north to
Angola and the Belgian Congo.

Genus CROTAPHOPELTIS Fitzinger

12a

1843. Crotaphopcltis Fitzinger, Syst. Eept., p. 27. Type by original

designation: Coronella rufescens Schlegel = Coronella hotam-

boeia Laurenti.
1927. DipsoglypliopJiLs Barbour & Amaral, Bull. Antivenin Inst. America,

1, p. 26. Type by original designation: Leptodira guineensis

Chabanaud.

Crotaphopeltis werneri (Boulenger)^^"

1897. Leptodira werneri Boulenger, Ann. Mag. Nat. Hist., (6) 19. p. 281:

Usambara Mountains, Tanganyika Territory.
1909. Dipsadomorphus reticulatus Werner, Jahresb. Ver. Nat. Wiirttem-

berg, 65, p. 55 : Tanga, Tanganyika Territorj'.
Range. Usambara Mountains and vicinity, Tanganyika Terri-
tory.

Crotaphopeltis hotamboeia kageleri Uthmoller

1939. Crotaphopeltis hotamboeia kageleri Uthmoller, Zool. Anz., 125,
p. 108: Sanya, Tanganyika Territorj'.

Range. Vicinity of Mount Kilimanjaro, Tanganyika Territory
(known only from the types).

Crotaphopeltis hotamboeia tomieri (Werner)
Sylvieoline White-lipped Snake
1908. Leptodira tomieri Werner, 1907, Sitzb. Akad. Wiss. Wien, 116.

Abt. 1, p. 1876 : Usambara Mountains, Tanganyika Territory.
Range. Virgin forests of the Usambara and Iluguru Moun-
tains, Tanganyika Territory, south to i\lisuku Mountains, north-
ern Nyasaland.

Crotaphopeltis hotamboeia hotamboeia (Laurenti)
Savanna White-lipped Snake
1768. Coronella hotamboeia Laurenti, Syn. Rept., p. 85: "India orien-
tali, ' ' i.e. Africa.

129 The included species present gradual stages of habitus from the slender
and attenuated arboreal forms like C. duchesnii guineensis to the moderately
stout terrestrial C. h. hotambO(.ia. A revision of the entire complex of this genus
in relation to Tclescopus, Dipsiidoha, Boiga and their included species, together
with a fresh evaluation of generic characters, is much needed. Piecemeal studies
result only in inconclusive reshuffling of forms.

130 1 suspect that the record of Dipsas obtusa from Usambara (Werner :1895).
repeateil by Tornier (1896) was based on this specimen.

272 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1768. Coronella virginica Laurenti, Syn. Rept., p. 86. No locality.

1789. Coluber nifescen^ Gmelin, Syst, Nat., ed. 13, 1. Part 3, p. 1094

(based on Seba, Thesaurus, 1, pi. xxxiii, fig. 6) : No locality.
1833. Coluber bicolor Leach in Bowdich, Miss. Ashantee, p. 493:

"Fantee," i.e. Fanti, Ashanti, Gold Coast.
1833. Ophis Heterurus Duvernoy, Ann. Sci. Nat. (Paris), 30, p. 9, pi. i,

fig. 2 : No locality.
1843. Dipsas Jiippocrepis Eeinhardt, Dansk. Vidensk. Selsk. Skrift, 10.

p. 251, pi. i, figs. 18-20 : Guinea.
1849. Dipsas inornatus A. Smith, Illus. Zool. S. Africa, Rept. App.,

p. 20: Kaffirland eastward of Cape Colony, i.e. Natal.
1860. Oxyropus melanocrotaphos Cope, Proc. Acad, Nat. Sci. Phila-
delphia, p. 2G0 : No locality.
1940. Tarbophis barnumhroivni Bogert, Bull. Amer. Mus. Nat. Hist., 77,

p. 66, fig. 9: Jigjiga, Ethiopia.
Range. Uganda ; Kenya Colony ; Tanganyika Territory ; Zanzi-
bar Island; indeed almost all Africa south of about 15° N.
(though unrecorded from French and British Somaliland).

Crotaphopeltis degeni (Boulenger)^"

1906. L.eptodira degeni Boulenger, Proc. Zool. Soc. London, p. 572, fig.

97: Entebbe, Uganda.

1907. Leptodira attarensis Werner, Sitzb. Akad. Wiss. Wien, 116, 1, p.

1875, pi. iii, fig. 6: Sennar and Sobat, Sudan.
Range. Southern Sudan and Uganda.

Genus CHAMAETORTUS Giinther

1864. Chamaetortus Giinther, Proc. Zool. Soc. London, p. 310. Type by
monotypy: C. aulicus Giinther.

Chamaetortus aulicus aulicus Giinther ^^-
Cross-barred Tree-Snake
1864. Cliamaetortus aulicus Giinther, Proc. Zool. Soc. London, p. 310:

Zambezi River.
Range. Kenya Colony; Tanganyika Territory and Zanzibar
Island south through Mozambique.

131 The relationship to C. h. hotambocia is obscure and a series is needed to
demonstrate that they really can be distinguished.

132 Trinomials are used as Mons. J. Guib6 assures me (30.V.195C) that the
young snake described in 1916 as C. a. ellenhergeri by Chabanaud appears to be
correctly assigned. As its type locality "Lambargne," Gabon, is well over 1500
miles from the nearest point where aulicus is known to occur, I suggest that
there may be an error in the typo locality, more especially as Mabuya striata
ellenbergeri was taken by the same collector on the Zambezi at Lealui, Northern
Rhodesia.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 273

Genus DISPHOLIDUS D

uvernov

1829. Bucephalus A. Smith (not Baer:1827), Zool. Journ., 4. p. 441.
Type by inonotypy (as other included species are synonyms) :
B. typus A. Smith.

1832. DLspholidus Duvernoy, Ann. Sci. Nat. (Paris), 26. p. 150. T3T)e by
monotypy: D. lalwndii Duvernoy = Bucephalus typus A. Smith.

1843. Dryomedusa Fitzinger, Syst. Eept., p. 26. Type by original desig-
nation: Dendrophis coluhrina Schlegel = Bucephulus typus A.
Smith.

DisphoHdus typus (Smith) "^

Boomslang or Back-fanged Tree-Snake
1829. Bucephalus typus A. Smith, Zool. Joum., 4. p. 441: Old Latakoo,

South Africa.^^*
1829. Buceplmlus jardinii A. Smith, Zool. Journ., 4, p. 442: South

Africa.
1829. Bucephalus gutturalis A. Smith, Zool. Journ., 4, p. 442: South

Africa.
1829. Bucephalus Bella A. Smith, Zool. Journ., 4. p. 442 : South Africa.
1832. DisphoHdus Lalandii Duvernoy, Ann. Sci. Nat. (Paris), 26, p.

150: Cape of Good Hope, i.e. South Africa.
1837. Dendrophis coluhrina Schlegel, Essai Phys. Serp., 2. p. 238, pi. ix,

figs. 14-16: Rondebosch, Cape Province, South Africa.
1841. Bucephalus viridis A. Smith, Illus. Zool. S. Africa, Kept., pi. iii :

Old Latakoo, South Africa.^^*
1841. Bucephalus capcnsis A. Smith, Illus. Zool. S. Africa, Eept., pis.

x-xiii: Cape Province, South Africa.
1849. Dendrophis pseudodipsas Bianconi, 1848, Nuovi Ann. Sci. Nat.,

(2) 10. p. 108, pi. iv, fig. 2; also 1850, Spec. Zool. Mosamb., p.

40, pi. iv, fig. 2: Mozambique.

133 That six of the synonyms should have been described from South Africa bv
Sir Andrew Smith, bears testimony to the astonishing variation in color, due iii
part to differences of age or sex. I have ijersoually taken four striking variants
in, or near, a single avenue of trees at Morogoro, Tanganyika Territory. The
all-black phase that occurs there, bears, both in color and scalation, so close a
resemblance to the aglyphous Thrasops jacksonii that one wonders if descent,
rather than convergence, is not the explanation.

The two races recently ilescribed by Laurent, are listed here because a paratype
of one form comes from Tanganyika Territory. Possibly they are recognizable but
their status rc(iiiires to lie studied in relation to the species as a whole, some-
thing I have not been able to do. The range of typus, as here understood, is as
extensive as that of Crotaphopeltis h. hotainboeia and nearly as great as that
of Bitis a. urietans, neither of which seems divisible to any appreciable extent.

134 Old Latakoo, or Lattakoo as it was sometimes spelled in the earlv days, it.
more ciirrectly rendered as Lithako. It was the principal kraal of the Batlaping
tribe, approximately 27° S., 24° E.. i.e. between Kuruman and Taungs, northern
Cape Province.

274 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1937. Thrasops jachsonii mossambicus Mertens, Abhand. Senckenberg.
Naturf. Ges., No. 435, p. 13: Cheringoma Farm, Inhaminga,
Mozambique.
1955. Wispholidus typus Tcivuensis Laurent, Revue Zool. Bot. Afr., 51.
p. 127: 3 from Uvira, 780 metres, Lake Tanganyika, Kivu
District, Belgian Euanda-Urundi.
1955. Wispholidus typus punctatus Laurent, Eevue Zool. Bot. Afr., 51,

p. 129: $ from Dundo, Angola.
Range. Uganda ; Kenya Colony ; Tanganyika Territory ; Pem-
ba, Zanzibar and Mafia Islands; indeed almost all Africa south
of about 15° N. (though unrecorded from Dahomey; Sierra Leone
and Gambia).

(Jenus THELOTORNIS Smith

1849. Thelotornis A. Smith, Illus. Zool. S. Africa, Rept., App. p. 19.

Type by monotypy: T. capcnsis A. Smith.
1859. Cladophis A. Dumeril, Arch. Mus. Hist. Nat. (Paris), 10. p. 204.

Type by monotypy: Leptophis kirtla>ndii Hallowell.

Thelotornis kirtlandii Idrtlondii (Hallowell)
Northern Vine-Snake ^^^
1844. Leptophis Kirtlandii Hallowell, Proc. Acad. Nat. Sci. Philadelphia,

p. 62 : Liberia.
1854. Oxyhelis Lecomtei Dumeril & Bibron, Erpet. Gen., 7. p. 821:

Gabon, French Congo.
1854. Tragops nifulns Dumeril & Bibron, Erpet. Gen., 7. p. 827: Senegal.
185(5. Oxyhelis violacea Fischer, Abhand. Nat. Ver. Hamburg, 3. p. 91,

pi. ii, fig. 7: Edina, Grand Bassa County, Liberia.
Range. Somalia, south through Uganda and Kenya Colony to
northern Tanganyika Territory (where it meets and merges with
the southern race) southAvest to central Angola, north and west
to Portuguese Guinea.

Thelotornis kirtlandii capensis Smith Southern Vine-Snake
1849. Thelotornis capensis A. Smith, Illus. Zool. S. Africa, Rept., App.,

p. 19: Kaffirland and the country towards Port Natal, i.e.

Durban, Natal.
1895. Thelotornis Kirtlandii var. mossamhicana Boeage, Herp. Angoht

Congo, p. 119: Manica, Mozambique (restricted).
Range. Southern Tanganyika Territory and Mafia Island,

133 A name slightly preferable to Blrrl-Snake, for, though its principal prey
may consist of birds and nestlings at certain seasons or in some localities, else-
where chameleons and other reptiles figure largely in its menu.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 275

south (except in area occupied by T. k. oatesii Giinther) to Natal
northwest to northern Southwest Africa and southern Angola
(where it meets and merges with the northern race).

Genus HEMIRHAGERRHIS Boettger

136

1893. Hemirhagerrhis Boettger, Zool. Anz., 16, p. 129. Type by mono-
typy : H. kelleri Boettger.

Hemirhagerrhis kelleri Boettger Stripe-bellied Snake

1893. Hemirhagerrhis kelleri Boettger, Zool. Anz., 16, p. 129: "Ah-

dallah," i.e. Abdulla, north of Webi Shebeli, Ethiopia.
1908. Amplarhinus taeniatus Sternfeld, Mitt. Zool. Mus. Berlin, 4. p.

244, fig. 3: Lamu Island, Kenya Colony.
Range. Ethiopia and British Somaliland, south through So-
malia to Mombasa and Voi, Kenya Colony.

Hemirhagerrhis nototaenia nototaenia ((liinther)
Eastern Spot-striped Snake
1864. Coronella nototaenia Giinther, Proe. Zool. Soc. London, p. 309,

pi. xxvi, fig. 1 : Eios de Sena, Zambezi, Mozambique.
1878. Ablabes Eildebrandtii Peters, Monatsb. Akad. Wiss. Berlin, p.

205, pi. ii, fig. 6: Kitui, Ukamba, Kenya Colony.
1901). Amplorhinus Giintheri Mocquard, Bull. Mus. Hist. Nat. (Paris),

12, p. 251: Lake Ngami, Bechuanaland Protectorate.
Range. Sudan and British Somaliland south through Kenya
Colony and Tanganyika Territory to Mozambique and Bechuana-
land. north through the Rhodesias to Belgian Cono-o.

Genus PSAMMOPHYLAX Fitzinger

137

1768. Cerastes Laurenti, Syn. Kept., p. 81. Type by subsequent designa
tion of rejervary:1923: Coluber rhombeatus Linnaeus. Type
by subsequent designation of Stejneger :1936: Cerastes candidus
Laurenti =: Coluber haje Linnaeus, i.e. Naja haje (Linnaeus).

136 Bogert (1940. Bull. Amer. Mus. Xat. Hist., 77, p. 75) considers Amplorhintig
A. Smith (1847. 111. Zool. S. Africa. Kept., text to pi. Ivii) must for the present
be regarded as a monotypic genus, with A. multimaciilatus Smith the only
species.

137 Technically Cerastes is correct, but in view of its application to the horned
vipers of North Africa for so long (I have over 100 citations in this sense
between 1880 and 1950), to use it for the more or less innocuous opisthoglyphous
grass-snakes of East and South Africa seems to imply a pedantic devotion to
legality. In view of the medical implications involved in anti-cerastes serum and
the frequency with which the name Cerastes has appeared in medical literature,
it should be re.iected, or Stejneger's solution of the difficulty accepted.

276 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1843. Psammophylax Fitzinger, Syst. Kept., p. 26. Type by original

designation: Coluber rhombeatus Linnaeus.
1847. T rimer orhinus A. Smith, Illus. Zool. S. Africa, Eept., text to pi.

Ivi. Type by monotypy: Coluber rhombeatus Linnaeus.

Psammophylax tritaeniatus multisqucanis ( Loveridge ) ^' "
Many-scaled Grass-Snake
1932. T rimer orhinus tritaeniatus multisquamis Loveridge, Proc. Biol.

Soc. Washington, 45, p. 84: Nairobi, Kenya Colony.
Range. Highlands of Ethiopia and Kenya Colony, south (in
highlands) to the Central Railway, Tanganyika Territory.

Psammophylax tritaeniatus variabilis Giinther ^■^''
Gray-l)e]lied Grass-Snake
1893. Psammophylax variabilis Giinther, Proc. Zool. Soc. London, 1892,

p. 5ri7, col. pi. XXXV : Shire Highlands, Nrasaland.
Range. Highlands of Tanganyika Territory (south of the Cen-
tral Railway), south (in highlands) to Mozambique and Nyasa-
land.

Psammophylax tritaeniatus tritaeniatus (^Giinther)
White-bellied Grass-Snake
1868. Rhageri'his tritaeniatus Giinther, Ann. Mag. Nat. Hist., (4) 1. p.

423, pi. xix, fig. H: Southeastern Africa.
1881. Coronella tritaenia Giinther, in Frank Gates, Matabeleland and
the Victoria Falls (London), App. 3, p. 329, pi. C: "South-
eastern Africa," i.e. Matabeleland, Southern Rhodesia.
Range. Dry savanna of Tanganyika Territory-, south to Natal,
west to Southwest Africa, north to Angola, east through south-
ern Belgian Congo to Lake Tanganyika.

Genus RHAMPHIOPHIS Peters

18.54. Rhamphiophis Peters, Monatsb, Akad. Wiss. Berlin, p. 624. Type

by monotypy: B. rostratus Peters.
1863. Dipsina Jan, Arch. Zool. Anat. Fisiol., 2, pp. 215, 313. Type by

monotypy: Coronella multimaculata A. Smith.

138 Early Kenya and Tanganyika records appeared as Rhagerrhis tritaeniata ,
my own first captures were erroneously recorded as Psammophis hrevirostrin.
Boulenger's (1896:649) record of Trimerorhinus tritaeniatus from "Uganda,"
like otliers from the same source, should read Kenya Colony.

139 The "Trimerorhinus rhombeatus" record of Sternfeld (1910) from Ukinga.
southern Tanganyika Territory, was almost certainly based on a misidentitiPii
P. t. variabilis, the form which is abundant throughout the southern highlands.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 277

Rhconphiophis acutus acutus (Giinther)
Southern Striped Beaked-Snake
1888. Psammophis acutus Giinther, Ann. Mag. Nat. Hist., (6) 1, p. 327,

pi. xix, fig. D: Pungo Ndongo, Angola.
Range. Southern Tanganyika Territory west through North-
ern Rhodesia to Angola, northeast through the Belgian Congo
to Belgian Kuanda-Urundi.

Rhconphiophis rubropunctatus (Fischer)
Red-spotted Beaked-Snake
1884. Dipsina ruhropunctaia Fischer, Jahrb. Hamburg. Wiss. Anst., 1,

p. 7, pi. i, fig. 3: Near Arusha, Tanganyika Territory.
Range. Sudan east to British Somaliland, south through So-
malia and eastern Kem'a Colony to northeastern Tanganyika
Territory, and {jide Moreau and Pakenham) Zanzibar Island.

Rhamphiophis oxyrhynchus rostratus Peters
Eastern Brown Beaked-Snake
1854. Pihumphiophis rostratus Peters, Monatsb. Akad. Wiss. Berlin, p.

624: Tete; Mesuril and Quitangonha, Mozambique.
1868. Fihagerrhis unguiculata Giinther, Ann. Mag. Nat. Hist., (4) \. p.

422, pi. xix, fig. G: Zanzibar.
1870. Coelopeltis porrectus Jan, Icon. Gen. Ophid., livr. 34, pi. ii, fig. 1:

Africa?
Range. Sudan east to Somalia, south through Kenya Colony ;
Tanganyika Territory and Zanzibar Island to Mozambique, west
through Nyasaland and Northern Rhodesia to southeastern
Belgian Congo.

Rhamphiophis oxyrhynchus oxyrhnchus (Reinhardt)
Western Brown Beaked-Snake
1843. Psammophis oxyrhynchus Reinhardt, Dansk. Vidensk, Selsk. Skrift.,

10. p. 244, pi. i, figs. 10-12: Guinea.
1929. Ehamphiophis connali Parker, Ann. Mag. Nat. Hist., (10) 4, p.

450: Accra, Gold Coast.
Range. Uganda, west through northern Belgian Congo to
Nigeria; Togo; Gold Coast; French Soudan.

Genus DROMOPHIS Peters "°

1843. Philodendros Fitzinger, Syst. Rept., p. 26. Type by original

140 Unfortunately this name is antedated by Philodendros, a most misleading
name for a terrestrial snake that bears a strong resemblance to Paammophia.
The sooner a ruling to reject Philodendros can be obtained, the better.

278 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

designation: Dendrophis praeornata Schlegel.
1846. Philodendrus Agassiz, Nomen. Zool., Index Univers., p. 285.

Emendation for Philodendros Fitzinger.
1869. Dromophis Peters, Monatsb. Akad. Wiss. Berlin, p. 447. Type by

monotypy: Dendrophis praeornata Schlegel.

Dromophis lineatus (Dumeril & Bibron)
Buff-striped Grass-Snake
1854. Dryophylax lineatus Dumeril & Bibron, Erpet. Gen., 7. p. 1124:
White Nile, Sudan.

1901. Psammopliis sibilans tumbensis Schenkel, Verb. Naturf. Ges. Basel,

13, p. 172: Tumbo Island, French Guinea.

1902. Psammopliis hrevirostris temporalis Werner, Verb. Zool. Bot. Ges.

Wien, 52. p. 335: Coja, Togo.
Range. Sudan and Uganda, south along Lake Tanganyika,
Tanganyika Territory to noilhern Nyasaland and Northern
Rhodesia, northwest through the Belgian Congo to Portuguese
Guinea (though as yet not recorded from the Gold Coast and
Sierra Leone ) .

Genus PSAMMOPHIS Boie"^

1819. Macrosoma Leach (not Hubner:1818), in Bowdich, Miss. Ashantec,
App., p. 493. Tj'pe by monotypy: Coluber elegans Shaw.

1826. Psammophis Boie, Isis von Oken, 19, col. 982. Type by monotypy:
Coluber sibilans Linnaeus.

1826. Psammophis Fitzinger, Neue Class. Rept., pp. 29, 30. Type accord-

ing to Boie: Coluber sibilams Linnaeus.

1827. Psa7nmophis Boie (part), Isis von Oken, 20, col. 521. Type by

specific designation: Cohiber sibilans Linnaeus.
1838. Taphrometopon Brandt, Bull. Acad. Sci. St. Petersburg, 3, p. 243.

Type by monotypy: Coluber (T.) Uneolatus Brandt.
1854. Chorisodon Dumeril & Bibron, Erpet. Gen., 7. p. 901. Type by

monotypy: C. sibiricum Dumeril & Biljron = Coluber Uneolatus

Brandt.
1854. Monodiastema G. Bibron, in Dumeril & Bibron, Erpet. Gen., 7,

p. 901. Type by monotypy: Chorisodon sibericum Dumeril &

Bibron = Coluber (Taphrometopon) Uneolatus Brandt.
1868. Phayrea Theobald, Cat. Rept. Asiatic Soc. Mus., p. 51. Type by

monotypy: P. isabellina Theobald = Coluber condanarits Mer-

rem.

141 Owing to some confusion as to authorship I have cited all three of tlic
earliest records ; Fitzinger attributes the name to Boie.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 279

1872. Amphiophis Boeage, Jorn. Sci. Lisboa, 4. p. 81. Type by mono-

typy: A. angolensis Boeage.
1892. Ehamphophis Boulenger, Zool. Eec. for 1891, p. 12. New name

for Amphiophis Boeage, preoecupied.
1924. Mike Werner, Sitzb. Akad. Wiss. Wien, 133, Abt. 1, p. 51. Type

by monotypy: M. elcgantissimus Werner = Coluber condanarus

Merrem.

Psommophis punctulcrtus trivirgotus Peters
Southern Speckled Sand-Snake
1878. Psavimophis punctulatus var. trivirgatus Peters, Monatsb, Akad.

Wiss. Berlin, p. 206 "Taita," i.e. Teita, Kenya Colony.
Range. British Somaliland and Somalia, south through
Uganda ( ?) and Kenya Colony to Arusha and Same, Tanganyika
Territory.

Psommophis sibilons sibilons (Linnaeus) Hissing Sand-Snake
1758. Coluber sibilans Linnaeus (part), Syst. Nat., ed. 10, 1, p. 222:
"Asia," (error).

1802. Coluber Gemmatus Shaw, Gen. Zool., 3, p. 535: No locality.

1803. Coluber moniliger Daudin, Hist. Nat. Kept., 7, p. 69: No locality.
1827. Coluber auritus Geoffroy Saint-Hilaire, Kept., in Descr. Egypte,

pp. 147, 151, pi. viii, fig. 4: Egypt.
1856. Psammophi^i irregularis Fischer, Abhand. Nat. Ver. Hamburg, 3.
p. 92: Peki, Gold Coast.

1881. Psammophis brevirostris Peters, Sitzb. Ges. Naturf. Freunde

Berlin, p. 89: Xa-Matlale, Mozambique.

1882. Psammophis sibilaiis var. mossambica Peters, Reise nach Mossam-

bique, 3, p. 122: Mozambique Island, Mozambique.
1882. Psammophis sibilant var. tettensis Peters, Reise nach Mossambique,

3, p. 122: Tete and Mozambique Island, Mozambique.
1884. Psammophis sibilans var. intermedius Fischer, Jahrb. Hamburg.

Wiss. Anst., 1, p. 14: Arusha, Tanganyika Territory.
1887. Psammophis sibilans var. leopardinus Boeage, Jorn. Sci. Lisboa,

11. p. 206: Catumbela and interior Mossamedes, Angola.
1908. Psammophis thomasi Gough, Ann. Transvaal Mus., 1, p. 30, fig. — :

Salisbury, Southern Rhodesia.
Range. In suitable savanna country from Egypt south through
Uganda; Kenya Colony; Tanganyika Territory and Zanzibar
Island to Natal ; northwest to Southwest Africa and on to
Mauretania (outside of forested areas occupied by P. s. phil-
lipsii) ; east to the Sudan.

280 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Psominophis subtaeniatus sudanensis Werner
Northern Stripe-bellied Sand-Snake
1919. Psainmophis suhtaeniattis var. sudanensis Werner, Denks. Akad.

Wiss. Wien, 96. p. 504: Kadugli, Sudan.
Range. Drier regions of the southern Sudan and southern
Ethiopia, south through Uganda; Kenj^a Colony; Tanganyika
Territory ; Zanzibar and Mafia Islands, to northern Mozambique
and Nyasaland."^

Psammophis biseriatus tangcmicus Loveridge
Western Link-marked Sand-Snake
1940. Psammophis biseriatus tanganicus Loveridge, Bull. Mus. Comp.

Zool., 87, p. 57: Mangasini, Usandawi, Tanganyika Territory.
Range. Southern Libya, southeast through the Sudan to
Eritrea, British Somaliland and Ethiopia (where it meets with
P. h. hiseriatus along the Ethiopian-Somali border north of the
Nogal River) ; south through Uganda to western Tanganyika
Territory.

Psammophis biseriatus biseriatus Peters
Eastern Link-marked Sand-Snake
1881. Psammophis biseriatus Peters, Sitzb. Ges. Naturf. Freunde Berlin,

p. 88: "Taita," i.e. Teita, Kenya Colony.
1913. Psammophis bitaeniatus (lapsus for biseriatus) Boettger, in
Voeltzkow, Eeise in Ost-Af rika, 3, p. 355 : Patta Island, Kenya
Colony.
Range. Somalia, south through eastern Kenya Colony to
northeastern Tanganyika Territory (where it meets with P. h.
tanganicus in the vicinity of Lake Manka).

Psammophis angolensis (Boeage) Pigmy Sand-Snake

1872. Amphiophis angolensis Socage, Jom. Sci. Lisboa, 4. p. 82: Dondo,

Cuanza River, Angola.
1877. Ablabes Homeyeri Peters, Monatsb. Akad. Wiss. Berlin, p. 620:

Pungo Ndongo, Angola.
Range. Tanganyika Territory and Zanzibar Island, south to
Mozambique, west through Nyasaland and Northern Rhodesia to
Angola and the Belgian Congo.

142 The typical form P. s. subtaeniatus Peters (with its synonyms P. moniliger
var. bilineaius Peters :1S67 ; P. locagii Boulenger :1895 ; and P. tratisvaalenais
Gough:1908), characterized by three labials entering the orbit, occurs in central
Mozambique and extends right across southern Africa south of the Zambezi.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 281

Genus CALAMELAPS Gunther"'

1849. Choristodon A. Smith (not Jonas :1844), Illus. Zool. S. Africa,
Kept., App., p. 18: Type by monotypy: C. concolor Smith.

1866. Calamelaps Giinther, Ann. Mag. Nat. Hist., (3) 18, p. 26. Type
by original designation: Calamaria unicolor Eeinhardt.

1947. Choristocalamus Witte & Laurent, Mem. Mus. Hist. Nat. Belgique,
29, p. 16. New name for Choristodon Smith (preoccupied).

Calamelaps unicolor unicolor (Reinhardt) "*
Seventeen-scaled Purple-Glossed Snake
1843. Calamaria, unicolor Eeinhardt, Dansk. Vidensk. Selsk. Skrift., 10,

p. 236, pi. i, figs. 1-3: Guinea.
1877. Atractaspis Eildebrandtii Peters, Monatsb. Akad. Wiss. Berlin, p.

616, pi. — , fig. 3: Zanzibar Coast, i.e. Tanganyika Territory.
1923. Calamelaps niangarae Schmidt, Bull. Amer. Mus. Nat. Hist., 49,

p. 117, fig. 12: Niangara, Belgian Congo.
Range. Uganda ; Kenya Colony and Tanganyika Territory,
west to Sierra Leone.

Calamelaps unicolor warreni Boulenger
Nineteen-scaled Purple-glossed Snake
1908. Calamelaps warreni Boulenger, Ann. Natal Mus., 1. pp. 230, 234,

fig. 3 : Kosi Bay, Zululand.
1915. Calamelaps mellandi Boulenger, Proc. Zool. Soc. London, p. 214:

Chirini Island, Lake Bangweulu, Northern Rhodesia.
Range. Kenya Colony and Tanganyika Territory, south
through Mozambique and the Rhodesias to the Transvaal and
Zululand, Natal.

Calamelaps unicolor polylepis Bocage

Twenty-one scaled Purple-glossed Snake
1873. Calamelaps polylepis Bocage, Jorn. Sci. Lisboa, 4, p. 216: Dondo,
Cuanza River, Angola.

i-*3 Under the name of Choristocalamus, Witte and Laurent (loc. cit.) would
revive Smith's monot.vpic genus Choristodon on the grounds that it has retained
an anterior temporal, a seventh upper labial, and posterior sublinguals — charac-
ters that seem inadequate to me.

On the other hand their action in synonymizing Rhinocalamus with Cala-
melaps has much to commend it, but the genotype - — which is the only species
known to me — occupies so intermediate a position in relation to certain other
genera that I have preferred to retain it until further work has been done on
the group.

144 Only 17-scaied snakes occur in West Africa north of the Equator ; in the
east they appear to be associated with the surviving montane or gallerv forests,
but occasional individuals (about one in twenty) occur at Llwale in southeastern
Tanganyika Territory where 19-scaled specimens predominate. For these the nam*-
C. u. warreni is retained — at least until its distribution is worked out in rela-
tion to that of C. u. polylepis which appears to have only 21 scales in Angola.

282 BU1.LETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1888. Calnmelaps miolepis Giinther, Anu. Mag. Nat. Hist., (6) 1, p. 323:

Cape Maclear, Lake Nyasa, Nyasaland.
Rcmge. Tanganyika Territory {fide a single old record from
Tukuyu), south to Nyasaland; Southern Rhodesia; Angola.

Genus RHINOCALAMUS Giinther

1888. Ehinocalatmis Giinther, Ann. Mag. Nat. Hist., (6) 1, p. 322. Type
by monotypy : ff. dimidiatus Giinther.

Rhinocolaxnus dimidiatus Giinther

1888. Bhino calamus dimidiatus Giinther, Ann. Mag. Nat. Hist., (6) 1,
p. 322, pi. xix, fig. C: Mpwapwa, Ugogo, Tanganyika Territory.

Range. Tanganyika Territory (known only from the type
locality).

Genus MICRELAPS Boettger

1880. Micrelaps Boettger, Ber. Senckenberg. Naturf. Ges., p. 136. Type

by monotypy : M. mixlleri Boettger.
1888. Elaposchema Mocquard, Mem. Cent. Soc. Philom. Paris, p. 122.

Type by monotypy: E. vaillanti Mocquard.

Micrelaps bicolorotus Sternfeld

1908. Micrelaps bicoloratus Sternfeld, Sitzb. Ges. Naturf. Freunde Ber-
lin, p. 93 : Kibwezi, Kenya Colony.
1908. BhinocMamus melcagris Sternfeld, Mitt. Zool. Mus. Berlin, 4, p.

244, fig. 4: Lamu Island, Kenya Colony.
Range. Kenya Colony and Tanganyika Territory (N.E. of
Lake Manyara).

Genus AMBLYODIPSAS Peters

1856. Amblyodipsas Peters, Monatsb. Akad. Wiss. Berlin, p. 592. Type
by monotypy: Calamaria microphthalma Bianconi.

Amblyodipsas katangensis ionidesi Loveridge

1951. Amblyodipsas hatangensis ionidesi Loveridge, Bull. Mus. Comp.
Zool., 106, p. 193: Tunduru, Southern Province, Tanganyika
I'erritory.
Range. Tanganyika Territory.

LOVERIDQE: E. AFRICAN REPTILES AND AMPHIBIANS 283

Genus MIODON Dumeril"'

1858. Microsoma Jan (not Macquart:1855), Revue Mag. Zool,, (2) 10.

p. 519. Type by monotypy: M. neuwiedii Jan.

1859. Miodon A. Dum6ril, Arch. Mus. Hist. Nat. (Paris), 10. p. 206.

Type by monotypy: Elapomorphus gahonensis A. Dum^ril.

1860. Urobelus Reinhardt, Vidensk. Meddel, Kjebenhavn, p. 229. Type

by monotypy : U. acanthias Reinhardt.

1902. Cynodontophis Werner, Verb. Zool. Bot. Ges. Wien, 52. p. 346.

Type by monotypy: C. aemulans Werner = Microsoma notatum
Peters.
1941. Melanocalamu^ Witte, Explor. Pare. Nat. Albert, Miss. G. F. de
Witte, Fasc. 33, p. 216. Type by monotypy: M. leopoldi Witte.

Miodon colloris collaris (Peters) Pale-collared Snake-eater
1881. Microsoma collare Peters, Sitzb. Ges. Naturf. Freunde Berlin, p.
148: Macange, "Cuango" i.e. Kwango, French Congo.

1887. Microsoma fulvicollis Mocquard, Bull. Soc. Philom. Paris, (7) 11.

p. 65 : Franceville, French Congo.

1888. Elapomorphus caeutiens Giinther, Ann. Mag. Nat. Hist., (6) 1, p.

323, pi. xix, fig. B: Cameroon Mountain, British Cameroon.
1910. Cynodontophis werneri L. Miiller, Abhand. Bayer Akad. Wiss.,

2 Kl., 24. p. 612: Cameroon.
Range. Southwestern Uganda, west to Cameroon and possibly
Nigeria.

NCodon collaris chxistyi Boulenger Eastern Snake-eater

1903. Miodon Christyi Boulenger, Ann. Mag. Nat. Hist., (7) 12. p. 354:

Uganda.
1923. Miodon unicolor Schmidt, Bull. Amer. Mus. Nat. Hist., 49, p. 119,

fig. 13 : Poko, Belgian Congo.
Range. Central Uganda, south to western Tanganyika Terri-
tory, west to eastern Belgian Congo.

1*5 It is impossible at this time to investigate the numerous changes proposed
by Witte and Laurent (1947, Mem. Mus. Kov. Hist. Nat., 29, pp. 58-89) but
two ^ 5 gahonensis (M.C.Z. 49727, 53737) from the Gold Coast (which ther
would perhaps assign to neuwiedi Jan) have 218 and 233 ventrals respectively,
while two $ $ (M.C.Z. 53738, etc.) have 247 and 259. This would appear to
effectively dispose of the alleged race M. g. schmidti Witte and Laurent from
the Congo. On zoogeographical grounds, together with size and color pattern, I
accept their separation of gahonensis from the collaris group, but not on the basis
of adult eye diameter — being at least twice its distance from the mouth in
gahonensis^ as opposed to less than twice its distance in collaris, fulvicollis, etc.
Our material fails to support this allegation for the lirst two collaris — taken at
random — had eye diameters that were included 1 Ms (M.C.Z. 42956 :Congo) to 2
or rather more than 2 times (M.C.Z. 9254 :Cameroon) their distance from the
border of the mouth — a very poor character indeed.

284 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Miodon collaris groueri Sternfeld
Central-Lake-Region Snake-eater
1908. Miodon Graueri Sternfeld, Sitzb. Ges. Naturf. Freuude Berlin,

p. 94: "Entetbe," i.e. Entebbe, Uganda."^
Rcoige. Central Uganda, southwest to eastern Belgian Congo
(L. Kivu).

Genus CHILORHINOPHIS Werner"'

1907. ChilorhinopMs Werner, Akad. Anz. Wien, 44, p. 479 (brief notice)
and 1908 (for 1907), Sitzb. Akad. Wiss. Wien, 116. Abt. 1,
p. 1881. Type by monotypy: C. butleri Werner.

1927. Parkerophis Barbour & Amaral, Bull. Antivenin Inst. America, 1.
p. 25. Type by original designation: Apostolepis gerardi Bou-
lenger.

Chilorhinophis gerardi tanganyikae Loveridge
Tanganyika Two-headed Snake
1951. Chilorhinophis gerardi tanganyikae Loveridge, Bull. Mus. Conip.
Zool., 108, p. 195: Nyarakolo, Lake Tanganyika, Northern Rho-
desia.
Range. Western Tangaujaka Territory (Ujiji), south to
Northern Rhodesia (Nyamkolo), west to southern Belgian Congo
(Lukonzolwa).

Chilorhinophis carpenteri liwalensis Loveridge
Liwale Two-headed Snake
1951. Chilorhinophis carpenteri liwalensis Loveridge, Bull. Mus. Comp.
Zool., 106, y. 196: Liwale, Southern Province, Tanganyika Terri-
tory.
Range. Southeastern Tanganyika Territory.

Genus APARALLACTUS Smith

1849. Aparallactus A. Smith, lUus. Zool. S. Africa, Kept., App., p. 15.

Type by monotypy: A. capensis Smith.
1849. Elapomorphus A. Smith (not of Wiegmann : in Fitzinger :1843),

^46 Entebbe may be viewed with suspicion for Capt. C. R. S. Pitman, wlio re-
sided at Entebbe for many years, failed to find it there and doubts its occurrence.
Allegedly Grauer, like some other collectors, failed to label his material immedi-
ately. The nearest point to Entebbe from which the Museum of Comparative
Zoology has c. graueri, is Kingani, 5000 feet, near Fort Portal (M.C.Z. 54737 :
coll. J. H. Blower, Esq. xi.l955).

147 The Amani, Usambara Mountains, record of the Sudanese butleri is believed
to have lieen based on a misidentification, possibly of an Aparallactus werneri.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 285

Illus. Zool. S. Africa, Kept., App. p. 16, footnote. Type by
monotypy: E. capensis Smith.
1854. Uriechis Peters, Monatsb. Akad. Wiss. Berlin, p. 623. Type by
subsequent designation : U. lunulatus Peters.

1859. Elapops Giinther, Ann. Mag. Nat, Hist., (3) 4. p. 161. Type by

monotypy : E. modestus Giinther.

1860. Periaspis Cope, Proe. Acad. Nat. Sci. Philadelphia, pp. 241, 266.

Type by monotypy: P. plumheatra Cope = Elapops modestus

Giinther.
1863. Cercocalamus Giinther, Ann. Mag. Nat. Hist., (3) 11. p. 21.

Type by monotypy: C. collaris Giinther = Aparallactus capensis

Smith.
1870. Metopophis Peters, Monatsb. Akad. Wiss. Berlin, p. 643, pi. i,

fig. 3. Type by monotypy: Uriechis lineatus Peters.
1917. Houleophvi Chabanaud, Bull. Mus. Hist. Nat. (Paris), 1916, 22. p.

379. Type by monotypy: R. ch.evalieri Chabanaud z= Uriechis

lineatus Peters.
1923. GuyoTtutrchia Angel, Bull. Mus. Hist. Nat. (Paris), 29, p. 348. Type

by monotypy: G. unicolor Angel = Elapops modestus Giinther.

Aparallactus modestus (Giinther)"^
Western-Forest Centipede-eater

1859. Elapops modestus Giinther, Ann. Mag. Nat. Hist., (3) 4. p. 161,

pi. iv, fig. C: West Africa.

1860. Periaspis plumheatra Cope, Proc. Acad. Nat. Sci. Philadelphia,

p. 242 : Liberia.

1896. Aparallactus houlengeri Werner, Verh. Zool. Bot. Ges. Wien, 46.

p. 363, pi. vi, figs. 6-6b : Cameroon.

1897. Aparallactus peraffinis Werner, Verh. Zool. Bot. Ges. Wien, 47.

p. 404, pi. ii, fig. 3 : Interior of Cameroon.
1897. Aparallactus ubangensis Boulenger, Ann. Mag. Nat. Hist., (6)

19. p. 279, fig. — : Zongo, Ubangi Rapids, Belgian Congo.
1901. Aparallactus flavitorques Boulenger, Ann. Mus. Congo, (1) 2,

p. 11, pi. iv, fig. 3: Lubue, Kasai, Belgian Congo.

148 Material of modestus in the Museum of Comparative Zoology lends no sup-
port whatever to the action of Witte and Laurent (1947, pp. 99, 103) in reviving
ubangensis as a race of modestus, to whose synonymy I had relegated it in 1944
(p. 187). The lower labials do not conform to their key and, taken from East to
West we find the parietal is in contact with the 5th upper labial only (1 side) or
.5th and 6th (5 sides) or 6th upper labial (2 sides) in Uganda snakes.
5th only (2 sides) on a ^ (M.C.Z. 53994) from Mayala, B. C.

parietal in contact with 6th upper labial (M.C.Z. 42944) at Makaia, B. C.
5th and 6th (4 ex.) or 6th upper labial (2 ex.) at Bitye, F. C.
oth and 6th (1 ex.) at Accra, G. C.

5th and 6th (1 ex.), or 6th upper labial (1 ex.) in i.iberia.

In other words, both m. modestus and m. ubangensis occur in Liberia ; Gold
Coast (or possibly not ni. modestus); French Cameroon; Belgian Congo and
Uganda.

286 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1902. Aparallactus dolloi Werner, Verh. Zool. Bot. Ges. Wien, 52, p.

346: Banzyville, Ubangi River, Belgian Congo.
1902. Aparallactus congicus Werner, Verh. Zool. Bot. Ges, Wien, 52.

p. 346 : Lingunda, Belgian Congo.
1907. Aparallactus Bat.esii Boulenger, Ann. Mag. Nat. Hist., (7) 19, p.

325: 5 miles inland from Kribi, French Cameroon.
1910. Aparallactus Cliristyi Boulenger, Ann. Mag. Nat. Hist., (8) 5,

p. 512: Mabira Forest, "Chagwe, " i.e. Kyagwe, Uganda.
1917. Aparallactus nigrocollaris Chabanaud, Bull. Mus. Hist. Nat.

(Paris), 1916, 22. p. 377, figs. 18-19: French Congo.
1917. Aparallactus nigrocollaris Roucheti Chabanaud, Bull. Mus. Hist.

Nat. (Paris), 1916, 22. p. 378, figs. 20-21: French Congo.

1923. Guyomarchia unicolor Angel, Bull. Mus. Hist. Nat. (Paris), 29. p.

348, figs. 1-4: (probably near Sangha) French Congo.

1924. Aparallactus Graueri Werner, Sitzb. Akad. Wiss. Wien, 133. Abt.

1, p. 42: Beni, Belgian Congo.
Range. Uganda, west to Sierra Leone.

Aparallactus lunulotus lunulotus (Peters)
Southeastern Plumbeous Centipede-eater
1854. Uriechis lunulotus Peters, Monatsb. Akad. Wiss. Berlin, p. 623:

Tete, Mozambique.
Range. Sudan and Belgian Congo, south to Uganda (Mt. Rom)
and Tanganyika Territory ; Mozambique ; Nyasaland ; the Rho-
desias and Transvaal.

Aparallactus liuiulatus concolor (Fischer)
Northeastern Plumbeous Centipede-eater
1884. Uriechis concolor Fischer, Jahrb. Hamburg. Wiss. Anst., I. p. 4,

pi. i, fig. 1 : Arusha, Tanganyika Territory.
1931. Aparallactus concolor boulengeri Scortecci (not Werner :1896),
Atti. Soc. Sci. Nat. Ital. (Milano), 70. p. 212: Villaggio Duca
degli Abruzzi and inland from Mogadiscio, Somalia.
Range. Sudan, east to Eritrea, south through eastern Kenya
Colony to northern Tanganyika Territory.

Aparallactus jacksonii jacksonii (Giinther)
Kilimanjaro Centipede-eater
1888. Uriechis Jacksonii Giinther, Ann. Mag. Nat. Hist., (6) 1, p. 325,
pi. xix, fig. E: Foot of Mount Kilimanjaro, Tanganyika Terri-
tory.
Range. Southern Ethiopia, south through Kenya Colony to
Tanganyika Territory."**

149 A race (A. j. oweni Loveridge) occurs at Torit in the southern Sudan.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 287

Aparollactus wemeri Boiilenger Usambara Centipede-eater
1895. Aparallactus Wemeri Boulenger, Ann. Mag. Nat. Hist., (6) 16<

p. 172: Usambara Mountains, Tanganyika Territory.
Range. Eastern Tanganyika Territory.

Aparallactus turner! Loveridge Malindi Centipede-eater

1935. Aparallactus turneri Loveridge, Bull. Mus. Comp. Zool., 79, p. 9:

Sokoki Forest, near Malindi, Kenya Colony.
Range. Coastal Kenya Colony (from Lamu to Mombasa).

Aparallactus guentheri Boulenger ^^° Zanzibar Centipede-eater
1895. Aparallactus guentheri Boulenger, Ann. Mag. Nat. Hist., (6) 16.
p. 172: "East and Central Africa" (omit Angola). Subse-
quently types were designated as from Zanzibar; Shire High-
lands and Lake Nyasa, Nyasaland.
1928. Aparallactus uluguruensis Barbour & Loveridge, Mem. Mus. Comp.
Zool., 50, p. 132: Nyange, Uluguru Mountains, Tanganyika
Territory.
Range. Coastal Kenya Colony; Tanganyika Territory and
"Zanzibar," south to Nyasaland.

Aparallactus capensis capensis Smith Cape Centipede-eater
1849. Aparallactus capensis A. Smith, Illus. Zool. S. Africa, Eept., App.,
p. 16: "Kaffirland eastward to Cape Colony," i.e. Natal, South
Africa.
1849. Elapomorphus capensis A. Smith, Illus. Zool. S. Africa, Rept. App.,
p. 16: "Kaffirland eastward of Cape Colony," i.e. Natal, South
Africa.
1863. Cercocalamus collaris Giinther, Ann. Mag. Nat. Hist., (3) 11.

p. 21, pi. iii, fig. A: "Central America" (error).
1895. Aparallactus punctatolineatus Boulenger, Ann. Mag. Nat. Hist.,

(6) 16. p. 173: Angola.
Range. Tanganyika Territory; Zanzibar and Mafia Islands:
south through Mozambique ; Nyasaland and the Rhodesias to
Cape Province (East London), northwest through Bechuanaland
to Angola.

Subfamily DASYPELTINAE
Genus DASYPELTIS Wagler'^'

150 A., guentheri was based on the ring-necked young, uluguruensis on the uni-
formly plumbeous adult.

151 statements to the contrary notwithstanding, the gender of Dasypeltis is
feminine. At the time this check list goes to press the genus is undergoing re-
visionary study by Carl Gans.

288 , BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1829. Anodon A. Smith (not of Oken:1815),i" Zool. Journ., 4, p. 443.

Type by monotypy: A. typiis Smith = Coluber scaber Linnaeus.

1830. Dasypeltis Wagler, Nat. Syst. Amphib., p. 178. Type by mono-

typy: Coluber scaber lavamsievis.
1830. Analcis Wagler, Nat. Syst. Amphib., p. 191, footnote 3. New
name for Anodon A. Smith (preoccupied).

1833. Rachiodon Jourdan, Le Temps (Paris:13.vi.l833), p. ?. New

name for Anodon A. Smith (preoccupied).

1834. Eaphiodon Jourdan (misprint) L'Institut (de la France: Paris)

2, p. 214, corrected to Rachiodon on p. 223.

1845. Deirodon Owen, Odontography; or, A Treatise on the Comparative

Anatomy of the Teeth . . . (London), p. 220. New name for
Anodon A. Smith (preoccupied).

1846. Dirodon Agassiz, Nomen. Zool., Index Univers., p. 127. Emenda-

tion for Deirodon Owen.

Dasypeltis scabra (Linnaeus) ^'"^ Common Egg-eater

1758. Coluber scaher Linnaeus, Syst. Nat., ed. 10, 1, p. 223: "in Indis, "

i.e. South Africa.i^*
1829. Anodon typus A. Smith, Zool. Journ., 4, p. 443: Near Capetown,

Cape Province, South Africa.

1853. Rachiodon abyssinus A. M. C. Dumeril {nom&n nudum), Mem.

Acad. Sci. (Paris), 23, p. 71: A casual reference to Cartin
(misspelled Quartin) Dillon's specimen from Ethiopia.

1854. Rachiodon Abyssinus Dumeril & Bibron, Erpet. Gen., 7, p. 496:

* ' Abyssinia, ' ' i.e. Ethiopia.

1864. Dasypeltis scaber var. capensis Peters, Monatsb. Akad. Wiss. Ber-
lin, p. 644, footnote: Cape of Good Hope, i.e. South Africa.

1864. Dasypeltis scaber var. mossambicus Peters, Monatsb. Akad. Wiss.
Berlin, p. 644, footnote: Boror and Tete, Mozambique.

1864. Dasypeltis scaber var. breviceps Peters, Monatsb. Akad. Wiss. Ber-
lin, p. 645, footnote: "Kaf&rland," i.e. South Africa.

1878. Dasypeltis lineolatris Peters, Monatsb. Akad. Wiss. Berlin, p. 206:
Kitui, Ukamba, Kenya Colony.

1912. Dasypeltis scabra var. atra Sternfeld, Wiss. Ergebn, Deutsch.
Zentral-Afrika-Exped. 1907-1908, 4, p. 272 : Virgin forest behind
boundary mountains on northwest shore of Lake Tanganyika,
Belgian Congo.

1954. Dasypeltis scabra loveridgei Mertens, Zool. Anz., 152. p. 213:

152 Suppressed in favor of Dasypeltis in 1956 (cf. Opiu. Declar. Inter. Comm.
Zool. Nomencl., 12, p. 241. Opinion 387).

153 Dr. Gans points out that the species scabra, whicli may possibly be subject
to some racial subdivision, includes the specimens that in 1942 I referred to "s.
scaber" and "s. pabnariim." It is on Dr. Gans' advice that medici and fasciata
are now treated as full species.

15* Cf. Flower, 1933, Proc. Zool. Soc. London, p. 818.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 289

Finkenstein Farm near Windhoek, Southwest Africa.
Range. Southern Arabia and Somalia, south to Natal, west to
the Cape, northwest to the Gold Coast, east through the southern
Sudan to Ethiopia.

Dasypeltis medici lamuensis Gans

1957. Dasypeltis medici lamuensis Gans, Breviora (Cambridge, Mass.),
No. 79, p. 1 : Lamu Island, Kenya Colony.

Range. Reddish laterite or gray alluvial soils of coastal Som-
alia, south through Kenya Colony to the vicinity of Kilimanjaro
on the frontier of Tanganyika Territory.

Dasypeltis medici medici (Bianconi)

1859. Dipsas Medici Bianconi, Mem. Accad. Sci. Bologua, 10, p. 501, pi.

xxvi (reprinted as p. 277 of Spec. Mossamb.) : Mozambique.
1868. Dasypeltis scaber var. fasciolata Peters, Monatsb. Akad. Wiss.

Berlin, p. 451 ; Zanzibar Island.
1888. Dasypeltis elongata Mocquard, Mem. Cent. Soc. Philom. Paris,

p. 131, pi. xii, fig. 2: Zanzibar Island.
1893. Dasypeltis scabra var. bianconii "Med." Boettger (lapsus for var.

medici Bianconi), Zool. Auz., 16. p. 133.
Range. Reddish laterite soils of the Kenya Colony — Tan-
ganyika Territory frontier (as far inland as Kilimanjaro) ; Zan-
zibar and Mafia Islands ; south to Mozambique and Nyasaland.

Dasypeltis fasciata (Smith) Western-Forest Egg-Eater

1844. Dipsas carinatus Hallowell (not of Sehlegel:1837), Proc. Acad.

Xat. Sci. Philadelphia, p. 119: "Africa," later (1857, loc. cit..

p. 69) amended to Liberia.
1849. Dasypeltis fasciaius A. Smith, Illus. Zool. S. Afi-ica, Eept., foot

note to pi. Ixxiii : Sierra Leone.
1859. Bachiodon siibfasciatus A. Dumeril (nomen nudum), Arch. Mus.

Hist. Nat. (Paris), 10, p. 198: casual mention of name.
18G3. Eachiodon. scaber Tar. sntbfasciatus Jan (nomen nudum), Elenco

Sist. Degli Ofidi, p. 100: Gold Coast.
1863. BacModon scaber var. %micolor Jan (part: ■nomen nudum), Elenco

Sist. Degli Ofidi, p. 106: Gold Coast.
1907. Dasypeltis macrops Boulenger, Ann. Mag. Nat. Hist., (7) 19, p.

324: Efulen, French Cameroon.
Range. Western Uganda, west through the Belgian Congo and
Cameroons to Gambia.

290 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Family ELAPIDAE
Geniis ELAPSOroEA Bocage

1866. Elapsoidea Boeage, Jorn. Sci. Lisboa, 1. pp. 50, 70. Type by
monotypy : E. giintherii Boeage.

1896. Elapechis Boulenger, Cat. Snakes Brit. Mus., 3, p. 358, footnote.

Substitute name for Elapsoidea Boeage.

EQopsoidea sundevallii giintherii Boeage Western Garter-Snake
1866, Elapsoidea Giintherii Boeage, Jorn. Sei. Lisboa, 1. pp. 50 ,70, pi. i,
figs. 3-3b: Bissau, Portuguese Guinea and Cabinda, Portuguese
Congo.

1897. Elapechis moebiusi Werner, Verb. Zool. Bot. Ges. Wien, 47. p.

400: Kete, Gold Coast.
Range. Northern and Western Tanganyika Territory, through
western Kenya Colony and Uganda, west to Senegal.

Elapsoidea sundevallii loveridgei Parker
Northeastern Garter-Snake
1949. Elapsoidea sundevallii loveridgei Parker, Zool. Verhand. Rijksmus,

Nat. Hist. Leiden, No. 6, p. 95 : Machakos, Kenya Colony.
Range. Central and eastern Kenya Colony.

Elapsoidea sundevallii nigra Giinther Eastern Garter-Snake
1888. Elapsoidea nigra Giinther, Ann. Mag. Nat. Hist., (6) 1, p. 322:
' ' Ushambola, Zanzibar, ' ' i.e. Usambara Mountains, Tanganyika
Territory.
Range. Virgin forests of the Usambara; Magrotto and Ulu-
guru Montains, eastern Tanganyika Territory.

Elapsoidea simdevallii decosteri Boulenger
Southeastern Garter-Snake
1888. Elapsoidea Decosteri Boulenger, Ann. Mag. Nat. Hist., (6) 2.

p. 141 : Delagoa Bay, Mozambique.
1895. Elapsoidea Boulengeri Boettger, Zool. Anz., 18, p. 62: Boroma,
Zambezi, Mozambique.

Range. Southern Province of Tanganyika Territory, south
through Mozambique, Nyasaland and the Rhodesias to the Trans-
vaal.

Genus BOULENGERINA Dollo'^'

1886. Boulengerina Dollo, Bull. Mus. Roy. Hist. Nat. Belgique, 4. p.

155 For Bogert's reasons for svnonymlzing Limnonaja, see Bogert, 1943, BuU.
Amer. Mus. Nat. Hist., 81. pp. 296, 300.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 291

159. Type by monotypy: B. storTusi Dollo.
1923. Limnonaja Schmidt, Bull. Amer. Mus. Nat. Hist., 49, p. 124. Type
by monotypy: Boulengerina christyi Boulenger.

Bonlengerina annulata stormsi Dollo Tanganyika Water-Cobra
1886. Boulengerina stormsi Dollo, Bull. Mus. Eoy. Hist. Nat. Belgique,

4. p. 160, fig. — : Lake Tanganyika, Belgian Congo.
Range. Lake Tanganyika and its shores off Tanganyika Terri-
tory and Belgian Congo. ^^^

Genus NAJA Laurenti

1768. Naja Laurenti, Syn. Kept., p. 90. Type by tautonomy: Coluber

naja Linnaeus. ^'^
1820. Naia Merrem, Vers. Syst. Amphib., pp. 15 (bis), 147, 190.

Emendation for Naja Laurenti (Naja on p. 15 (primus)).
1830. Uraeus Wagler, Nat. Syst. Amphib., p. 173. Type by monotypy:

Coluber haje Linnaeus.

1830. Aspis Wagler (not Laurenti: 1768), Nat. Syst. Amphib., p. 173.

Type by monotypy: Coluber naja Linnaeus.

1831. Tomyris Eichwald, Zool. Spec. Eossiae Poloninae, Part 3, p. 171.

Type by monotypy : T. oxiana Eichwald ^ Coluber naja Lin-
naeus.

Naja haje haje (Linnaeus) Egyptian Cobra

1758. Coluber haje Linnaeus, Syst. Nat., ed. 10, 1. p. 225: Lower Egypt.

1768. Cerastes candidus Laurenti, Syn. Kept., p. 83: Libya.

1789. Coluber Candidissimus Lacep^de, Hist. Nat. Quad. Ovip. Serpens,

2. Table Methodique, p. 86, text p. 118: Libya.
1854. Naja Int.ermixta Dumeril & Bibron, Erpet. Gen., 7, p. 1299:

Based on pi. xix of A. Smith, 1849, so South Africa.
1854. Naja haje var. annulifera Peters, Monatsb. Akad. Wiss. Berlin,

p. 624: Tete, Mozambique.
1873. Naja haje var. viridis Peters, Monatsb. Akad. Wiss. Berlin, p. 411,

pi. i, fig. 1 : West Africa.

156 The locality data for the two British Museum specimens allegedly from
Nyasaland had best be regarded with suspicion pending confirmation. Mr. J. C.
Battersby informs me (16.i.57) that they were collected by Surgeon H. M.
Hanschell, R.N., for the London School of Tropical Medicine which, in 1926.
turned them over to the British Museum.

157 Stejneger (1936, Copeia, p. 140) states that Laurenti's six included species
are all synonyms of Coluber naja Linnaeus. Prior action by Boie (1826. Isis von
Oken, col. 981) in designating Vipera haemachatea Latreille as type of Naja is
inadmissable.

Included in the synonymy by some authors is Lepidon Swainson (1839,
2, p. 345). However, no type was designated, the name simply appearing in a
key between Naia and Elaps with the scant description : Head broad, neck not
expansive. // East African this might apply to Pseudohaje or Elapsoidea.

292 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Range. Spanish Morocco east to Egypt, south through Uganda,
Kenya Colony, Tanganyika Territory and Zanzibar Island. In
fact almost all Africa except for the area of N. h. harotseensis
Angel (with 15 midbody scale-rows) known only from Lealui,
Northern Rhodesia ; and tlie area occupied by N. h. anchietae
Bocage (with 17 midbody scale-rows) which ranges through the
Rhodesias, Bechuanaland, Southwest Africa and Angola. An-
other race occurs in Arabia.

Naja nigricollis pallida Boulenger Pink-or-red Spitting-Cobra
1896. Naia nigricollis var. pallida Boulenger, Cat. Snakes Brit. Mus., 3.
p. 379: Inland from Berbera, British Somaliland (restricted by
Bogert:1942), also Lake Eudolf.
Range. On red murram soils from northern Ghana and Sudan
east to British Somaliland, south through Kenj^a Colony (Voi
to Kibwezi) to Tanganyika Territory (vicinity of Longido Moun-
tain) .

Naja nigricollis nigricollis Reinhardt Common Spitting-Cobra
1843. Naja nigricollis Reinhardt, Danks. Vidensk. Selsk. Skrift., 10,

p. 369, pi. iii, figs. 5 and 7: Guinea.
1854. Naja mossamhica Peters, Monatsb. Akad. Wiss. Berlin, p. 625:

Sena and Tete, Mossambique.
1893. Naja nigricollis var. crawshayi Giinther, Proc. Zool. Soc. London,

p. 620: Lake Mweru, Northern Rhodesia.
189.J. N'aja nigricollis var. occidontalis Bocage, Herp. Angola Congo,
p. 135; Dondo; Huilla; Humbe; Quisange; and Quilengues,
Angola.
1895. Naja nigricollis var. melanolcuca Bocage (not of Hallowell:1857),
Herp. Angola Congo, p. 136: Caeonda and Catumbella, Angola.
1955. Naja nigricollis atriceps Laurent, Revue Zool. Bot. Afr., 51, p.
135 : Mugera, 1450 metres, Ruyigi Territory, Urundi, Belgian
Ruanda-Urundi.^^^
Range. Savanna areas of almost all Africa south of 25° N.
(except where occupied by recognizable races), thus including
Uganda ; Kenya Colony ; Tanganyika Territory ; Pemba and
Zanzibar Islands.

158 Specimens from Mangasini and other localities in Tanganyika Territory are
listed among the paratypes. Like most cobras nigricollis is very variable as
regards color, and sevei'al striking variants may occur in a single locality. Many
races not listed heie have been described. An overall study of the genus is
much to be desired.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 293

Naja melanoleuca Ilallowell Blaek-and-White Cobra ^'®

1857. Xaia haie var. melanoleuca Hallowell, Proc. Acad. Nat. Sci. Phila-

delphia, p. 61 : Gabon, French Congo.
1885. Naja haje var. leucosticta Fischer, Jahrb. Hamburg. "Wiss. Anst.,

2, p. 115, pi. iv, fig. 11 : Cameroon and Ogooue Kiver, French

Congo.
1955. ^Naja melanoleuca sutfulva Laurent, Eevue Zool. Bot. Afr., 51,

p. 132: Lwiro, 1850 metres, Kabare Territory, Kivu, Belgian

Congo.
Range. Forested areas of much of Africa south of 15° N. in
the "West, to the Equator in the East, including Uganda, Kenya
Colony, Tanganyika Territory and Zanzibar Island.

Genus PSEUDOHAJE Gunther

1858. Pseudohaje Giinther, Cat. Snakes Brit. Mus., p. 222. Type by

monotypy : P. nigra Giinther.

Pseudohaje goldii (Boulenger) Black Forest-Cobra

1S95. Xaia goldii Boulenger, Ann. Mag. Nat. Hist., (6) 16, p. 34: near

Asaba, Nigeria.
1896. NaAa yahoma Moequard, C.-E. Ill Congres Internat. Zool., Leyde,
1895, p. 233: Yakoma, Abiras District, northern Belgian Congo.
Range. Forested areas of Uganda, southwest to Southwest
Africa, northwest to Nigeria.

Genus DENDROASPIS Schlegel

1848.^^° Dendroaspis Schlegel, Versl. Zool. Gen. Amsterdam, p. 5. Type

by monotypy: Elaps jamesoni Traill.
1852. Dinophis Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p. 203.

Type by monotypy: D. hammondii Hallowell = Leptophis

viridis Hallowell.

1855. Dendroechis Fischer, Programm Realschule, Johann. Hamburg, pp.

20, 68. Type by monotypy: D. reticulata Fischer = Leptophis
viridis Hallowell.

1856. Dendraspis A. Dum^ril (not of Fitzinger:1843), Revue Mag. Zool.,

(2) 8, p. 558. Emendation for Dendroaspis Schlegel.

159 The few specimens I have taken in the southeast (Mikindani, T. T. ;
Nyasaland) were a variegated brown, not black as at Kaimosi, K. C. Whether
the brown cobras really occupy a well-defined range so that Laurent's name might
be applied to them, is not clear. Material in the Museum of Comparative Zoology
suggests there Is no such division In scale-counts between southeast and west as
is shown by Laurent's data.

160 Date uncertain, but communicated by Schlegel to a meeting held on March
13, 1848 (fide Brongersma, 1936, Zool. Meded., 19, p. 136).

294 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Dendroospis iamesoni koiinosae Loveridge
Eastern Jameson's Mamba
1936. Dendraspis jainesoni kaimosae Loveridge, Proc. Biol. Soc. Wash-
ington, 49. p. 64: Kaimosi, Kakamega, Kenya Colony.
Range. Western Kenya Colony, west through Uganda to
Belgian Ruanda-Urundi and the eastern Belgian Congo.

Dendroaspis angusticeps (Smith) Common Green Mamba
1849. Naia angusticeps A. Smith, lUus. Zool. S. Africa, Rept., pi. Ixx:

Natal, South Africa.
1865. Dendraspis intermedins Giinther, Ann. Mag. Nat. Hist., (3) 15, p.

97 : Zambezi Eiver, Mozambique.
1907. Dendraspis sjosiedti Lonnberg, ia Sjostedt, Wiss. Ergeb. Schwed.
Zool. Exped. KUimandjaro, Meru umgeb Massaisteppen, No. 4,
p. 17, pi. i, fig. 2: "Kibonoto, " i.e. Kibongoto, Mount Kiliman-
jaro, Tanganyika Territory.
Range. Kenya Colony; Tanganyika Territory and Zanzibar
Island, south through Mozambique, Nyasaland and the Rhodesias
to Natal.'"

Dendroaspis polylepis antinorii Peters'""
Northern Brown-Mamba
1873. Dendraspis antinorii Peters, Monatsb. Akad. Wiss. Berlin, p. 411,

pi. i, fig. 2: "Ansaba," i.e. Anseba, Eritrea.
Range. Eritrea, south through Ethiopia and British Somali-
land to Mount Moroto, northeastern Uganda, and Murri, northern
Kenya Colony.

Dendroaspis polylepis polylepis Giinther
Southern Brown-Mamba
1864. Dendraspis polylepis Giinther, Proc. Zool. Soc. London, p. 310:

Zambezi River, Mozambique.
1907. Dendraspis mamba Gough, Zool. Anz., 32, p. 454: White River

Settlement, near Nelspruit, Lydenburg District, Transvaal.
Range. Southern Kenya Colony and Tanganyika Territory,
south through Mozambique ; Nyasaland and the Rhodesias to
Natal, west to Southwest Africa, northeast through Angola to the
eastern Belgian Congo.

161 This is the correct distribution, confused for a century by the inclusion of
records pertaining to D. p. polylepis, etc.

162 For discussion on the validity of this race, see Parker (1949, Zool. Verb.
Rijksmus. Nat. Hist. Leiden, No. 6, p. 98). This, and typical polylepis are the
snakes commonly called "Black Mambas," though they are not black ; the young
are grey green, somewhat darker than the vivid fresh green of young and adult
angusticeps.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 295

Family HYDROPHIIDAE

*

Genus PELAMIS Daudin

1803. Pelajnis Daudin (part), Hist. Nat. Rept., 7, p. 361. Type by desig-
nation of Fitzinger:1843 : P. bicolor Daudin =: Anguis platvra
Linnaeus. ^^^

1816. Pelamys Oken, Lehrb. Naturg., 3. Part 2, pp. vi, xv, 279. New

name for Pelamis Daudin (part) with type by monotypy: Anguis
platuro (sic) Linnaeus.

1817. Ophincctes Rafinesque, Amer. Monthly Mag., 1, p, 432. Type by

present designation: 0. luteus Rafinesque.'^®*
1848. Elaphrodytes Gistel, Naturg. Thierr., p. ix. New name for

Hydnis Schneider {fide Stejneger:1907. Not seen.).
1910. Pelamydrus Stejneger, Proc. U. S. Nat. Mus., 38, p. 111. Type by

monotypy: Anguis platura Linnaeus.

Pelomis ploturus (Linnaeus) Parti-colored Sea-Snake

1766. Anguis platura Linnaeus, Syst. Nat., ed. 12, 1, p. 351: No locality.
1799. Hydrus Bicolor Schneider, Hist. Amphib., 1, p. 242 (based on

Scba, 1735, Thesaurus 2, pi. Ixxvii, fig. 2): Mexico?
1817. Pelamis sclineideri Rafinesque (new name for Hydrus bicolor

Schneider, var.), Amer. Monthly Mag., 1, p. 432.
1817. Ophinectes luteus Rafinesque, Amer. Monthly Mag., 1. p. 433: No

locality.^®^
1837. Hydrophis pelamis Sehlegel (new name for Anguis platura Lin-
naeus), Essai Phys. Serp., 1, p. 508.
1842. Pelamis ornata Gray, Zool. Misc., p. 60: Borneo.
1854. Pelamis bicolor var. variegata Dumeril & Bibron, Erpet. Gen., 7.

p. 1337: Celebes.
1854. Pelamis bicolor var. sinuata Dumeril & Bibron, Erpet. Gen., 7,

p. 1338 : No locality.
1856. Hydrophis (Pelamis) bicolor var. alternans Fischer (new name

for variegata Dumeril & Bibron), Abhand. Nat. Ver. Hamburg,

3, p. 63.
1872. Hydrophis bicolor var. macidata Jan, Icon. Gen. Ophid., livr. 40,

pi. iii: Indian Ocean.

163 For lengthy discussion on rejection cf Hydrus Schneider by Malcolm Smith
(1926, Monogr. Sea-Snakes, p. 116), see H. M. Smith and B. H. Taylor (1945.
U. S. Nat. Mus. Bull. 187, p. 176, footnote).

164 Rafinesque states that his new genus differs from Pelamis in its "carinated
or angular abdomen," and the species luteus is characterized as "entirely yellow."
It, and doubtless some or all of the other "species" of Ophinectes he describes,
is apparently but a variant of the highly variable Anguis platura Linnaeus.

165 As pointed out in the preceding footnote to the genus Ophinectes, possibly
several other "species" attributed to this genus are also synonyms that should
be added to the above list.

296 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1921. Hydrus platurus var. subohscurus Wall, Snakes of Ceylon, p. 422

(name for var. C of Boulenger:1896) : Ceylon.
1921. Hydrus platurus var. pallidus Wall, Snakes of Ceylon, p. 422

(name for var. D of Boulenger :1896) : Travancore, India.
1955. Pelamis platura vara, hrunnea ; fasciata; leucostriata ; neurileu-
cura; neuricatenata ; tricolor Deraniyagala, Colored Atlas of
some Vertebrates of Ceylon (Colombo), 3, p. 80; no precise
localities mentioned, merely "Ceylon waters."
Range. Indian and other tropical oceans as far as the coast
of Mexico in the Pacific Ocean. Occasionally temperate seas such
as those of South Africa and around New Zealand in the southern
hemisphere ; as far north as the Russian Coast Province in the
northern hemisphere.^*'*'

Family VIPERIDAE

Genus ATRACTASPIS Smith
1849. Atractaspis A. Smith, Illus. Zool. S. Africa, Rept., footnote to text

of pi. Ixxi. Type by monotypy: A. bibronii Smith.
1854. Brachycranion Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p.

99. Type by monotypy: B. corpulentum Hallowell.
1862. Eurystephus Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 337.

Type by monotypy: Elaps irregularis Reinhardt.
1895. Clothelaps Cope, Trans. Amer. Philos. Soc, 18. p. 211. Type by

present designation; Atractaspis congica Peters.
1906. Melanelaps Wall, Journ. Bombay Nat. Hist. Soc, 17. p. 27. Type

by monotypy: M. mcphersoni Wall = Atractaspis andersonii

Boulenger.

Atractaspis microlepidota microlepidota Glinther^®^
Small-scaled Burrowing-Adder

166 1 am unaware of any actual specimens of platurus from Kenya Colony,
Tanganyika Territory or Zanzibar Island : reported "sea-snakes" invariably
proved "to be highly colored muraenid eels. However, P. platurus has been taken
along the Red Sea coast of Somalia ; in the Seychelles and Madagascar, while
•"It is known from all parts of the coast of East Africa, where it is not uncom-
mon south of the equator," according to Malcolm Smith (1943, Fauna British
India, 3, Serpentes, p. 477).

167 Laurent, in his revision of the genus Atractaspia (1950:10) regards fallow
as an eastern race of microlepidota, whose typical form he restricts to the south-
ern Sudan,, but I am far from happy regarding the numerous races of microlepi-
dota postulated by Laurent. In the first place the type locality of fallax is not
in Tanganyika Territory as assumed by Laurent, but "drei Tagereisen von
Kiriama" and that is clearly shown on the map of Von der Decken's itinerary
(1S69, Reisen in Ost-Africa, 1, part 1, map iii ; also cf. part 2, 1871, pp. 64, 435)
as about halfway between Bura and Mombasa. In later years (1897) Tornier
corrupted the spelling to "Kircamo."

When adequate series are available it is interesting to note the extensive range
of variation of typical microlepidota. For example 14 ^ ^ from Torit, Sudan,
have a ventral range of 216-239; subcaudals 25-29; 9 99, also from Torit,
have 233-242 ventrals ; 21-24 subcaudals. The sexes and extremes have been
carefully checked in these snakes (M.C.Z. 53535-53557).

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 297

1866. Atractaspis microlepidota Giinther, Ann. Mag. Nat. Hist., (3) 18.

p. 29, pi. vii, fig. 3: "Probably Weat Africa" (error).
1866. Atractaspis fallax Peters, Monatsb. Akad. Wiss. Berlin, p. 890:

"Kiriame" corrected to "Kiriama" = Giriama, Kenya Colony.
1913. Atractaspis phillipsi Barbour, Proc. Biol. Soc. Washington, 26,

p. 148: Singa, Sennar Province, Sudan.
Range. Southern Sudan, east through Ethiopia to Somalia,
south to Kenya Colony.^^*

Atractaspis irregularis bipostocularis Boulenger^®'
Eastern Variable Burrowing-Adder
1905. Atractaspis bipostocularis Boulenger, Ann. Mag. Nat. Hist., (7)

15, p. 190: Fort Hall near Mount Kenya, Kenya Colony.
1908. Atractaspis conradsi Sternfeld, Sitzb. Ges. Naturf. Freunde Ber-
lin, p. 9-1: Ukerewe Island, Lake Victoria, Tanganyika Territory.
1930. Atractaspis schoutedeni "Witte, Eevue Zool. Bot. Afr., 19, p. 224,

figs. 1-3: "N'Goma," i.e. Goma, Lake Kivu, Belgian Congo.
1934. Atractaspis bahaulti Angel, Bull. Soc. Zool. France, 59, p. 169:

Kadjudju, Kivu region, Belgian Congo.
1945. Atractaspis irregularis loveridgei Laurent, Eevue Zool. Bot. Afr.,

38, p. 322: Bunia, Ituri region, Belgian Congo.
Range. Kenya Colony and northwest Tanganyika Territory,
west through Uganda and Belgian Ruanda-Urundi to the eastern
Belgian Congo.

^o"-

Atractaspis oterrima Giinther

Western-Forest Burrowing-x\dder
1863. Atractaspis aterrima Giinther, Ann. Mag. Nat. Hist., (3) 12,
p. 363 : West Africa.

it>s Unknown from Uganda, for Lorlwar, though west of Lake Riulolf is in Kenya
Colony (not Uganda as stated by Parlter :1949 :109) ; the unique $ with 37
midbody scale-rows and 245 ventrals, allegedly from "Lake Tanganyilca" accord-
ing to Sir Jolin Kirk who sent it to the British Museum, is scarcely assignable to
this form.

169 Proceeding from east to west across Africa there is a gradual, but definite,
increase in scale-rows from 21 (Fort Hall) to 25-27 (i. irregularis at Ganta,
Liberia). A. bipostocularis was founded on a juvenile with two postoculars (no
second specimen has been taken in fifty years). I regard this condition as
aberrant, for In our series of 2o m. microlepidota from Torit, Sudan, two $ J

(M.C.Z. 53554-5) have two postoculars on the right side of the head but the
normal single scale on the left. If this view is accepted, then the name
bipotitocnlaris should take precedence over conradsi (which was also founded on
an aberrant individual as I demonstrated by collecting normal "irregularis" on
Ukerewe Island. Apart from Laurent's Eritrean i. angeli, and his i. uelensis
from the Congo, which are both apparently recognizable by high ventral counts,
the j^uestion arises as to how many forms one should recognize — 21-23, 23-25.
25-2 (, or just two, viz. 21-23 with some overlap of 25 In Uganda and eastern
Congo, and 25-27 for the western i. irregularis (inc. parkeri Laurent, if the
subcaudal counts are not found to overlap with more material).

298 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Range. Virgin forest of Uluguru Mountains, Tanganyika Ter-
ritory, northwest to Uganda, west to Portuguese Guinea.

Atractaspis bibronii rostrata Giinther^'"
Zanzibar Bibron's Burrowing-Adder
1868. Atractaspis rostrata Giinther, Ann. Mag. Nat. Hist., (4) 1, p. 429,

pi. xix, fig. 1 : Zanzibar.
1901. Atractaspis Icatangae Boulenger, Ann. Mus. Congo, (1) 2, p. 13,

pi. V, fig. 2: Lofoi, Katanga, Belgian Congo.
1901. Atractaspis coarti Boulenger, Ann. Mus. Congo, (1) 2, p. 14, pi. v,

fig. 3 : Albertville, Belgian Congo.
Range. Kenya Colony and Tanganyika Territory (Zanzibar
coast; possibly Zanzibar Island), south to Mozambique (north of
the Zambezi), west through Nyasaland and Northern Rhodesia
to southern Belgian Congo.

Genus CAUSUS AVagler

1830. Causus Wagler, Nat. Syst. Amphib., p. 172. Type by monotypy

(as Naja V-nigrum Boie is a synonym): Sepedon rhombeata

Lichtenstein.
1842. Distidhuriis Hallowell, Journ. Acad. Nat. Sei. Philadelphia, 8, p.

337. Type by monotypy: D. viaculatiis Hallowell = Sepedon

rhombeata Lichtenstein.
1862. Eeterophis Peters, Monatsb. Akad. Wiss. Berlin, p. 276. Type

by monotypy: H. resimus Peters.
1882. Dinodipsas Peters, Sitzb. Akad. Wiss. Berlin, p. 893. Type by

monotypy : D. angidifera Peters = Aspidelaps lichtensteinii Jan.

Cousus rhonxbeatus (Lichtenstein)^" Rhombic Night- Adder
1823. Sepedon rhombeata Lichtenstein, Verz. Doubl. Mus. Zool. Berlin,

p. 106 : No locality.
1827. Naja Col(uber) v nigrum Cuv. {sic) Boie, Isis von Oken, 20, col.

557: Africa.
1842. Bistichurus Maculatus Hallowell, Journ. Acad. Nat. Sci. Phila-
delphia, 8, p. 337, pi. xix: Liberia.

170 To this form all East African records of hibrotiii (to which can be added
Tornier's misidentiflcations of '■irregularis" from Tanga, etc.) are referable.
Laurent has shown that rostrata is distinguishable by normally having 23 mid-
body scale-rows (rarely 21 or 22), whereas h. bibronii A. Smith — with a range
south of the Zambezi to Angola — has 21 scale-rows (rarely 23).

171 I am not prepared to say that this common reptile, which (except for
Eritrea, Southwest Africa, and the offshore islands) has been reported from every
political unit in Africa south of 20° N., does not break up into some races. I
would point out, however, that small populations of uniformly colored, an'l
presumably striped individuals also, are scattered over much of the range and
occur together with normally marlied specimens. They do not seem to be any
more entitled to racial recognition than Vipera dorsalis Gray does in Britain.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 299

1905. Cajisus rhomheatus var. hilirifiatus Boulenger, Ann. Mag. Nat.

Hist., (7) 16, p. 114: Between Benguela and Bihe, Angola.
1912. Causus rhomheatus var. taeniata Sternfeld, Wiss. Ergebn. Deutscb.
Zentral-Afrika-Exped. 1907-1908, 4. p. 276: No. locality (but
Sehubotz coll., .so probaljiy between Lakes Victoria and Kivu).
1955. Caucus lineatus Laurent, Eevue Zool. Bot. Afr., 51, p. 136: Serani
Research Station, Kundelungu Plateau, Kasenga Territory,
Upper Katanga, Belgian Congo.
Range. Sudan east to British Soinaliland, south through
Uganda, Kenya Colony and Tanganyika Territory to Natal,
northwest to Angola and French West Africa (Mauretania and
French Soudan).

Causus resimus (Peters) Velvety-green Night- Adder

1862. Heterophis resimus Peters, Monatsb. Akad. Wiss. Berlin, p. 277.

pi. — , fig. 4: Jebel Ghule, Sennar, Sudan.
1888. Causus Jacksonii Giinther, Ann. Mag. Nat. Hist., (6) 1, p. 331:
Lamu, Kenya Colony; Laie Tanganyika, Tanganyika Terri-
tory.
1893. Causus 7iasalis Stejneger, Proc. U. S. Nat. Mus., 16, p. 735: Tana

River, Kenya Colony; Cunga, Angola; also West Africa.
1895. Causus resimus var. angolensis Bocage, Herp. Angola Congo, p

148: Rio Dande and eight other localities in Angola.
Range. Sudan, east to British Somaliland, south through
Uganda; Kenya Colony and Tanganj'ika Territory to Mozam-
bique, west tlirough Northern Rhodesia to eastern Belgian Congo
and Belgian Ruanda-Urundi.

Causus defilippii (Jan) Snouted Night- Adder

1862. H {.eterodon) Be Filippii Jan, Arch. Zool. Anat. Fisiol., 2. p. 225:

Africa.

1863. Causus {Eeteropliis) rostratus Giinther, Ann. Mag. Nat. Hist., (3)

12, p. 363: Ugogo, Tanganyika Territory.
Range. Coastal Colony ; Tanganyika Territory and Zanzibar
Island (at Mtende), south through Mozambique; Nyasaland and
the Rhodesias to Natal.

Cavisus lichtensteinii (Jan)

1859. Aspidclaps Lichtensteinii Jan, Revue Mag. Zool., p. 511: Gold

Coast.
1882. Dinodipsas angulifera Peters, Sitzb. Akad. Wiss. Berlin, p. 893,

pi. XV : "Laguna, near Puerto Cabello, Venezuela" (error).
Range. Western Kenya Colony and Uganda, west to Spanish
(luinea ; French and British Cameroons ; Nigeria ; Liberia.

300 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus VIPERA Laurenti

1768. Fipera Laurenti (part), Syn. Kept., p. 99. Type by designation

of Fitzinger:1843 : Vipera redii Gmelin r= Coluber francisci redi

Laurenti = Cohiier aspis Linnaeus.
1816. Bems Okeu, Lehrb. Naturg., 3, Part 2, p. 234. Type by tautonomy:

Coluber berus Linnaeus.
1820. Pelias Merrem, Vers. Syst. Amphib., p. 148. Type by monotypy:

Coluber berus Linnaeus.
1822. Chersea Fleming, Phil. Zool., 2. p. 295. Type by monotypy:

' ' Cerastes vulgaris " ? rr Vip.era vulgaris Latreille = Coluber

aspis Linnaeus.
1834. ETtinaspis Bonaparte, Icon. Fauna. Ital., Anfibi, part. 42. Type by

monotypy : Coluber ammodytes Linnaeus.

1842. Daboia Gray, Zool. Misc., p. 69. Type by subsequent designation:

Vipera elegans Daudin = Coluber russellii Shaw.

1843. Ehinechis Fitzinger, Syst. Eept., p. 28. Type by original designa-

tion: Coluber ammodytes Linnaeus.

1843. Chersophis Fitzinger, Syst. Kept., p. 28. Type by original designa-

tion: Vipera elegans Daudin = Coluber russellii Shaw.

1844. Echidnoides Mauduyt, Herpetol Vienne, p. 29. Type by original

designation: E. trilamina Millet =: Coluber berus Linnaeus.

1!)27. Mesocoronis Eeuss, Zool. Anz., 71. p. 216. Type by original desig-
nation : .V. c.oronis Eeuss = Vipera berus var. bosniensis Boett-
ger.

1927. Teleovipera Eeuss, Zool. Anz., 73. p. 125. Type by original desig
nation : Coluber ammodytes Linnaeus.

1927. Acridophaga Eeuss, Zool. Anz., 73. p. 126. Type by original
designation: Pelias ursinii Bonaparte.

1927. Mesovipcra Eeuss, Zool. Anz., 73. p. 126. Type by original desig-
nation: Coluber aspis Linnaeus.

1927. Macrovipera Eeuss, Zool. Anz., 73. p. 126. Type by original desig-
nation: Coluber lebetinus Linnaeus.

1927. Mesohorinis Eeuss, Naturforseher, 4. p. 129. Misprint for Meso-
coronis Eeuss.

1929. Latastea Eeuss, Wochenschr. Terrar. Aquar.-Kunde, 26. p. 64.
Type by original designation: Vipera latastei Bosca.

]!i29. Tzarevscya Eeuss, Wochenschr. Terrar. Aquar.-Kunde, 26. p. 64.
Type by original designation: Mesocoronis {Tzarevscya) tigrina
Eeuss =^ Vipera haznalcovi Nikolski.

19?.;". Latasteopara Eeuss, Nachr.-Blatt. Aquar.-Terrar.-Ver., p. 216.
Type by original designation : Vipera hugyi Schinz.

Vipera hindii Boulenger Montane Viper

1910. Vipera hindii Boulenger, Ann. Mag. Nat. Hist., (8) 5. p. 513:

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 301

Fort Hall, 4000 feet, Kenya Colony."^
Range. Kinangop and Aberdare Mountains, Kenya Colony.

Vipera supercilioris Peters Lowland Viper

1854. Vipera superciliaris Peters, Monatsb. Akad. Wiss. Berlin, p. 625:
Mainland opposite Querimba Island, Cape Delgado, Mozam-
bique.
Range. Mwaya, Lake Nyasa, Tanganyika Territory and Mo-
zambique.

Genus BITIS Gray

1768. Colra Laurenti,"^ Syn. Kept., p. 103. Type by designation of
ritzinger:1843: Colvher atropos Linnaeus.

1820. Echidna Merrem (part; not Forster :1788), Vers. Syst. Amphib., p.
150. Type by designation of Fitzinger :1843: E. arietans Mer-
rem.

1842. Clotho Gray (not Faujas de Saint-Fond: 1808), Zool. Misc., p. 69.
Type by tautonomy: Cohra clotho Laurenti.

1842. Bitis Gray, Zool. Misc., p. 69. Type by tautonomy: Coluber bitis
(sic: error for bitin) Bonnaterre = Cobra lachesis Laurenti.

1845. Calechidna Tschudi, Fauna Peruana, Herp., p. 60. Type by mono-
typy: Echidna ocellata Tschudi = Coluber atropos Linnaeus.

Bitis worthingtoni Parker Kenya Horned Viper

1932. Bitis worthingtoni Parker, Journ. Linn. Soc. London, Zool., 38,

p. 221 : Lake Naivasha, Kenya Colony.
Range. Uplands of Kenya Colony.

Bitis arietans arietans (Merrem) ^'^ Puff Adder

1768. Cobra lachesis Laurenti, Syn. Eept., p. 104: No locality.

1768. Cobra clotho Laurenti, Syn. Kept., p. 104: "Ceylon and Cuba"

(error).
1789. Cioluber) Bitin Bonnaterre, Tabl. Encycl. Method. Ungues Nat.,
Ophiol., p. 22: "Ceylon" (error).

172 One wonders whether the type actually came from Fort Hall, or was merely
sent by Dr. S. L. Hinde when stationed there.

173 Suppressed in favor of Bitis in 1945 (cf. Opin. Declar. Inter. Comm. Zool.
Nomencl., 3, pp. 77-89 : Opinion 188).

174 Actually the correct name is lachesis, which has been used less than a
dozen times since it was revived by Stejneger. As it has long been associated
generically with the Tropical American pit-vipers, the sooner it is officially
suppressed the better. Between 1849 and 1949 Bitis arietans has been con-
sistently used over 250 times in the literature. The English name, which has
come to stay, is also somewhat unfortunate for, while the burrowing-adders and
night-adders are all oviparous, the "puff adder" is a truly viviparous "viper."

302 BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY

1798. Coluber intumescens Donndorf, Zool. Beytrage Linn. Natur., 3.

p. 209: (not seen).
1802. Vipera severa Sonnini & Latreille (part), Hist. Nat. Kept., 3.

p. 335, pi. — , fig. 1: "Japan" (error).
1820. Vipera {Echidna) arietans Merrem, Vers. Syst. Aniphib., p. 152:

Cape of Good Hope, i.e. South Africa.
1822. Vipera inflata Burchell, Travels S. Africa, 1. p. 469: "Cape

Colony," i.e. Cape Province, South Africa.
1829. Vipera brachyura Cuvier, Kegne Animal, ed. 2, 2. p. 90: No

locality.
1842. Clotho lateristriga Gray, Zool. Misc., p. 69: Gambia.
Range. Africa (outside forested areas) from southern Morocco
east to Arabia; south (except for an undefined area centering
about British Somaliland where B. a. somalica Parker occurs)
through Uganda; Kenya Colony; Tanganyika Territory (and
possibly Zanzibar on the basis of a single specimen collected by
Kirk, though not necessarily on the island) to Natal, west
through Cape Province, northwest to Morocco.

Bitis arietans somalica Parker

1949. Bitia lackcsis somalicu Parker, Zool. Verhand. Eijksmus. Nat. Hist.

Leiden, No. 6, p. 101: Bohodle, 2100 feet, British Somaliland.
Range. British Somaliland, south to northern Kenya Colony.

Bitis gobonica gabonica (Dumeril & Bibron)
Central African Gaboon Viper ^"
1854. Eohidna Gabonica Dumeril & Bibron, Erpet. Gen., 7. p. 1428, pi.

Ixxx b: Gabon, French Congo.
Range. Southern Sudan and Uganda ; eastern Tanganyika
Territory ^"'^ south to Mozambique ; west through Northern Rho-
desia to Angola; north and West to Togoland where (or in the
Gold Coast) it meets with the race B. g. rhinoceros Schlegel
which ranges westward to French Guinea.

*t>^

Bitis nasicornis (Shaw) Nose-horned Viper

1802. Coluber Nasicornis Shaw, Nat. Miscell., 3. pi. xciv: Interior of
Africa (from tlie master of a Guinea vessel).

175 Also known as the River-Jack : the name Rhinoceros Viper, though properly
pertaining to the West African race of Gaboon Viper, is best aliandoned as it
has been so often applied to the Nose-horned Vi])er.

1T6 "Zanzibar" was included by me in 1924 on the basis of Sternfeld's 1910
listing. Doubtless his record, like that of Boulenger in 1896, referred to the
opposite coast (now Tanganyika) as it is unknown on the island today. Stern-
feld also lists gabonica from Damaraland. i.e. Southwest Africa, and Stevenson-
Ilamiltou includes it in the herpetofauna of Kruger I'ark, Transvaal. Both are
omitted from the above range pending contirmation.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 303

1854. Vipera Eexacera Dum^ril & Bibron, Erpet. Gen., 7. p. 1416, pi.
Ixxviii h, fig. 2 : No locality.

Range. Southern Sudan; western Kenya Colony; Uganda;
southwest through Belgian Ruanda-Urundi and the Belgian
Congo to Angola; north and west to French Guinea (but un-
recorded from Togo and Sierra Leone).

Genus ECHIS Merrem

1820, Echis Merrem (part). Vers. Syst. Amphib., p. 149. Type by
designation of Boie (1826) or Fitzinger (1843) : Pseudoboa
carinata Schneider.

1849. Toxicoa Gray. Cat. Snakes Brit. Mus., p. 29. Type by subsequent
selection: Echis arenicola Boie := Scythale pyramidum Geoffrey.

Echis ccorinotus pyramiduin (Geoflfroy)
Egyptian Saw-scaled Viper
1827. Scythulc pyramidum Geoffrey Saint-Hilaire, Kept., in Descr.

Egypte, p. 152, pi. vii, fig. 1: Egypt.
1827. Echis arenicola Boie, Isis von Oken, 20. col. 558: North Africa.
1834. Echis pavo Eeuss, Mus. Senckenberg., 1. p. 157: Egypt.
1834. Echis varia Eeuss, Mus. Senckenberg., 1, p. 160: Ethiopia.
1837. Fipcra echis Schlegel, Essai Phys. Serp., 2. p. 583. pi. xxi, figs.

10-11: Africa; Arabia; Bengal, India.
Range. Northern India, west through Persia and Arabia to
North Africa, south in arid regions to northern Kenya Colony ;
Uganda and Gold Coast. ^''

n'

Genus ATHERIS Cope

1849. Chloroechis Bonaparte, Proc. Zool. Soe. London, p. 145, footnote.
Type by tautonomy: Vipera chloroechis Schlegel.^'*

1862. Aiheris Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 337. Type by

monotypy: Vipera chloroechis Sclilegel.

1863. Poecilostolus Giinther, Ann. Mag. Nat. Hist., (3) 11. p. 25. Type

liy monotypy: P. hurtonii Giinther = Echis sqiiatxic/er Hal
lowell.

177 Possibly the Gold Coast form is separable. E. c. carinatus Schneider occurs
in a limited area around Madras, India.

178 Chloroechis was rejected by Boulenger on the grounds that it was inade-
(|uately described. However, Bonaparte refers to the green color of this true, but
arboreal, viper from Ashanti, and his evident intention was to erect a genus
whose type by tautonomy was Vipera chlnroechis Schlegel.

304 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Atheris squamiger (Hallowell) Green Bush-Viper^'®

1854. Echis squainigera Hallowell, Proc. Acad. Nat. Sci. Philadelphia,
p. 193 : Near Gabon Eiver, French Congo.

1863. Poecilostolus Biirtonii Giinther, Ann. Mag. Nat. Hist. (3), 11, p.

25: Cameroon.

1864. Atheris polylepis Peters, Monatsb. Akad. Wiss. Berlin, p. 642:

Liberia.
1885. Atheris Lucani Rochebrune, Bull. Soc. Philom. Paris (7), 9. p. 89:

Landana, Cabinda.
1885. Atheris proximus Rochebrune, Bull, Soc. Philom. Paris (7), 9,

p. 90: Bissarie, Casamance River district, Senegal.
1887. Atheris anisolepis Moequard, Bull. Soc. Philom. Paris (7) 11. p.

89 : " Alim Leketi, ' ' presumably Alima Leketi River, French

Congo.

1887. Atheris laeviccps Boettger, Zool. Anz., 10, p. 651: Povo Netonna,

near Banana, Belgian Congo.

1888. Atheris suhocularis Fischer, Jahrb. Hamburg. Wiss. Anst., 5, p. 5,

pis. i, fig. 1, and ii, fig. 11 : Cameroon.
1955. Atheris hispida Laurent, Revue Zool. Bot. Afr., 51, p. 138:

Lutungura, Kivu District, Belgian Congo.
Range. Western Kenya Colony and Uganda; west through
Belgian and French Congo to Cameroons and Fernando Po;
south to Angola.

Atheris nitschei nitschei Tornier Great Lakes Bush-Viper

1902. Atheris nitschei Tornier, Zool. Jahrb. Syst., 15, p. 589, fig. — :

Mpororo Swamp, Tanganyika Territory.
1906. Atheris woosnami Boulenger, Ann. Mag. Nat. Hist., (7) 18, p. 37:

Mubuku Valley, Euwenzori Mountains, Uganda.
Range. Uganda ; western Tanganyika Territory ; Belgian
Ruanda-Urundi ; Belgian Congo.

Atheris nitschei rungweensis Bogert
Rungwe Mountain Bush-Viper
1940. Atktris nitschei rungweensis Bogert, Bull. Amer. Mus. Nat. Hist.,
77, p. 104, fig. 18: Rungwe Mountain, Tanganyika Territory.
Range. Southwest Tanganyika Territory ; Misuku IMountains,
northwest Nyasaland and Northern Rhodesia.

1'9 Though commonly referred to as "tree-vipers," so far as my experience goes
these snakes are usually found in bushes or shrubbery at the forest-edge. Pos-
sibly trionomials should be applied to this form if it can be demonstrated that
chloroechis (Schlegel) is but a western race ranging from the Gold Coast to
Liberia.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 305

Athens cerotophorus Werner Usambara Mountain Bush-Viper
1895. Atheris ccratophorus Werner, Verb. Zool. Bot. Ges. Wien, 45,
p. 194, pi. V, fig. 1: Usambara Mountains, Tanganyika Terri
tory.

Range. Usambara Mountains, Tanganyika Territory .^^"

Atheris barbouri Loveridge Uzungwe Mountain Bush-Viper
1930. Atheris barbouri Loveridge, Proe. New England Zool. Club, 11,

p. 107: Dabaga, Uzungwe Mountains, Tanganyika Territory.
Range. T'zungwe and Ukinga Mountains, Tanganyika Terri-
tory.

Class AMPHIBIA

Order GYMNOPHIONA^'^

Family CAECILIIDAE^'-

Genus SCHISTOMETOPUM Parker

1941. Schistometopum Parker, Ann. Mag. Nat. Hist.. (11) 7, p. 17,
jfig. 4. Type by original designation: Dermophis gregorii Bou-
lenger.

Schistometopum gregorii (Boulenger)^*''
Mud-dwelling Caecilian
1894. Dermophis gregorii Boulenger, Proe. Zool. iSoc. London, p. G4tt,
pi. .\1, fig. 4: Ngatana, Tana River, Kenya Colony.

1912. Boulengerula denhardti Xieden, Sitzb. Ges. Naturf. Freunde Ber-

lin, p. 199: Tana Eegion, Kenya Colony.

1913. Bdellophis unicolor Boettger, in Voeltzkow, Reise in Ostafrika, 3,

p. 353, pi. xxiii, fig. 18: Lake Peecatoni, near Witu, Kenya
Colony.

Range. Coastal Kenya Colony, south to Wami River near
Bagamoyo. and Rufigi River near Kilindi, Tanganyika Territory.

Genus BOULENGERULA Tornier

1896. Boxdengcrula Tornier, Die Kriechthiere Deutsch-Ost-Afrikas, p.
164. Type by monotypy: B. boulengeri Tornier.

ISO The Togo and Gold Coast records are rejected pending definite capture.
181 APODA of r.nulenger, 1882, Cat. Batr. Gradientia s. Caudata and Batrachia
Apoda ill the Coll. Brit. Mus., p. 88.

'>^S2 Hypogrophis guentheri Boulenger, 1SS2, described as from "Zanzibar," has
been showu by Parker. 1941, to be a synon.vm of H. rontrata (Cuvier) of the Sp.v-
chelles. It is consequently omitted from this list.

1S3 The reasons for amalgamating these three "species," following the collectiou
of topotypes of all three, will be found in Lo%eridge, 1936, Bull. Mus. Comp.
Zool., 79, pp. 373-:>78.

306 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Boulengerula taitanus Loveridge Taita Mountains Caecilian
1935. Boulengerula taitanus Loveridge, Bull. Mus. Comp. Zool., 79. p. 16:
Mount Mbololo at 4800 feet, Taita Mountains, Kenya Colony.
Range. Taita Mountains, Kenya Colony.

Boulengerula changamwensis Loveridge
Cliangamwe Lowland Caecilian
1932. Boulengerula cliangamwensis Loveridge, Bull. Mus. Comp. Zool.,
72. p. 381 : Changamwe at 192 feet, near Mombasa, Kenya
Colony
Range. Kenya Colony (known only from the type locality).

Boulengerula boulengeri Tornier
Usambara Bluish-Gray Caecilian
1896. Boulengerula boulengeri Tornier, Die Kriechthiere Deutsch-Ost-
Afrikas, p. 16-i: Usambara Mountains, Tanganyika Territory.
Range. Usambara and Magrotto Mountains, Tanganyika Ter-
ritory.

Boulengerula uluguruensis Barbour & Loveridge
Uluguru Pink Caecilian
1928. Boulengerula uluguruensis Barbour & Loveridge, Mem. Mus. Comp.
Zool., 50. p. 183: Vituri, 2000 feet, Uluguru Mountains, Tan-
ganyika Territory.
Range. T^higuru Mountains, Tanganyika Territory.

Genus SCOLECOMORPHUS Boulenger

1883. Scolecomorplius Boulenger, Ann. Mag. Nat. Hist., (5) 11, p. 48.

Type by nionotypy : S. kirkii Boulenger.
1895. Bdellophis Boulenger, Proc. Zool. Soe. London, p. 412. Type by

monotypy : B. vittatus Boulenger.

Scolecomorphus vittatus (Boulenger)

1895. Bdellophis vittatus Boulenger, Proc. Zool. Soc. London, p. 412, pi.

xxiv, fig. 4: Usambara Mountains, Tanganyika Territory.
Range. Usambara; Magrotto and Uluguru Mountains, Tan-
ganyika Territory.

Scolecomorphus attenuatus Barbour & Loveridge

1928. Scolccomorplnis attenuatus Barbour & Loveridge, Mem. Mus. Comp.

Zool., 50, p. 181: Nyingwa, 7500 feet, Uluguru Mountains,

Tanganyika Territory.
Range. THuguru Mountains, Tanganyika Territory.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 307

Scolecomorphus kirkii uluguruensis Barbour & Loveridge
1928. Scolecomorphus uIugurKensis Barbour & Loveridge, Mem. Mus.
Comp Zool, 50, p. 180: Nyingwa, 7500 feet, Uluguru Mountains.
Tanganyika Territory.

Range. Uluguru IMountaius, Tanganyika Territory.

Scolecomorphus kirkii kirkii Boulenger

1883. Scolecomorphus Kirl-ii Boulenger, Ann. Mag. Nat. Hist., (5) 11,
p. 48: "Lake Tanganyika?"

Range. Highlands of .southwestern Tanganyika Territory;"^
Nyasaland and Northern Rhodesia ( Nyamkolo :/ic?e Pitman).

Order SALIENTIA

Suborder OPISTHOCOELA

Family PIPIDAE^^'

Genus XENOPUS AVagler

1827. Xenopus Wagler, Isis von Oken, 20, col. 726. Type by monotypy:

X. boiei Wagler = Bufo laevi-s Daudin.
1829. Dactylethra Cuvier, Regne Animal, ed. 2, 2, p. 107. Type by

monotypy: "crapaud lisse de Daudin" = Bufo laevis Daudin.
1864. Silurana Gray, Ann. Mag. Nat. Hist., (3) 14. p. 315. Type by

monotypy: S. tropicalis Gray.

Xenopus clivii Peracea Eritrean Smooth Clawed-Prog

1898. Xenopus Clivii Peracea, Boll. Mus. Zool. Anat. Comp. Univ.

Torino, 13. No. 321, p. 3: Adi Caie and Saganeiti, Eritrea.
Range. Eritrea, south through Ethiopia to Lake Rudolf,

Kenya Colony.

Xenopus laevis borealis Parker Kenya Smooth Clawed-Frog
1936. Xenopus laevis borealis Parker, Ann. Mag. Nat. Hist., (10) 18,

p. 596: Mount Marsabit, Kenya Colony.
Range. Kenj^a Colony.

Xenopus laevis victorianus Ahl

Tanganyika Smooth Clawed-Frog

1.S4 I regard Nieden's record of Bdellophis vittatus from Ubena as a misidenti-
lied kirkii for I personally collected this specie.s on the Mutindi-Njombe road, in
addition to a large series from Nyasaland.

1H5 Formerly referred to the suborder AGLOSSA, the African tongueless frogs
are separated by some authors as XENOPIDAE. Besides PIPIDAp; the OPIS-
THOCOELA includes only the DISCOGLOSSIDAE.

308 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1924. Xenopus victorianus Ahl, Zool. Anz., 60, p. 370: Busisi, Smith

Sound, Lake Victoria, Tanganyika Territory.
Range. Uganda ; Kenya Colony and western Tanganyika Ter-
ritory, west to Belgian Congo.

Xenopus laevis bunyoniensis Loveridge
Ruanda Smooth Clawed-Frog
1932. Xenopus laevis hunyoniensis Loveridge, Proc. Biol. Soc. Washing-
ton, 45, p. 114: Bufundi, Lake Bunyoni, Uganda,
Range. Highland lakes of southwest Uganda ; Belgian Ruanda
Urundi and Belgian Congo.

Xenopus laevis petersii Bocage
Angola Smooth Clawed-Frog
1895. Xenopus Petersii Bocage, Herp. Angola Congo, p. 187: Anibaca ;
Bcnguela; Caconda; Catumbela; Chiniboto; Dondo; Huila;
Quibula; Quindumbo and Sao Salvador, Angola.
1927. Xenopus poiceri Jlavntt, Ree. Albany Mus., 3, p. 413, pi. xxiv, fig.

3 : Victoria Falls, Zambezi, Northern Rhodesia.
Range. Southwestern Tanganyika Territory, southwest
through Northern Rhodesia to Angola.

Xenopus muelleri (Peters) Miiller's Smooth Clawed-Frog
1844. Dactylethra Muelleri Peters, Ber. Akad. Wiss. Berlin, p. 37:

Mozambique (restricted to Tete, Zambezi, by Loveridge).
Range. Southern Sudan (at Gondokoro) ; Uganda; Kenj^a
Colon}- ; Tanganyika Territory ; Zanzibar and Mafia Islands :
Mozambique soutli to Transvaal {fide Mertens) ; w^est through
Nyasaland and Northern Rhodesia to Angola ; north and west
in arid areas to Dahomey {fide Chabanaud).

Suborder PROCOELA
Family BUFONIDAE

Genus BUFO Laurenti^'"

1758. Bufo A. J. Roesel von Rosenhof, Hist. Nat. Ranarum, p. 69, pi.

xxiv. Type by present designation : B. aquaticus von Rosenhof

186 Bufo of von Rosenhof appears to have been ignored as pre-Linnaean and
not consistently binomial. Hylaplcsla Schlegel, 1826 (which antedates Boie,
1827) has been relegated to the synonymy of Dendrobates by Davis (1935, Zool.
Ser. Field Mus. Nat. Hist., 20, p. 88), whose genus Cacophryne (Type: Hi/l(i
borbonica Tschudi) is a synonym of Leptophryne Fitzinger, Syst. Rept. p. 32.
Type by original designation : Bm/o cruentatiis Tschudi. Phryniscus Wiegmann.
1834, whose type by monotypy is P. nigricans Wiegmann = Atelopua stelzneri
Weyenberg. is also one of the ATELOPODIDAB. f^cutiger Theobald, 1868, was
removed from the synonymy by Boulenger.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 309

equals and antedates B. calamita Laurenti.
1768. Bufo Laurenti (part), Syn. Eept., p. 25. Type by tautonomy: B.
vulgaris Laurenti = Bona bufo Linnaeus.

1824. Oxyrhynchus Spix, Anim. Spec. Testud. Eana. Brasiliam, p. 49.
Type by monotypy: 0. naricus Spix = Eana typhonm Lin-
naeus.

1828. Chaunu^ Wagler, Isis von Oken, 21, col. 744. Type by monotypy:
C. marmoratus Wagler (inc. C. glohulosiis Wagler) both =
Bufo granulosus Spix.

1838. Osilophu^ "Cuv." Tschudi, Class. Batr. Kept., p. 89. Type by
monotypy: " Osilophus typJionius Cuv. " = Eana typhonw
Linnaeus.

1843. Phrynoidis Fitziuger, Syst. Eept., p. 32. Type by original designa-
tion: Bufo asper "Kuhl, " i.e. Gravenhorst.

1843. Fhrynomorphus Fitzinger, Syst. Eept., p. 32. Type by original
designation: Bufo leschenaultii Dumeril & Bibron =: Bufo
guttaius Schneider.

1843. Bocidophryne Fitzinger, Syst. Eept., p. 32. Type by original
designation : Bufo agiia Daudin = Eana marina Linnaeus.

1843. PeJtophryne Fitzinger, Syst. Eept., p. 32. Type by original desig
nation: Biifo peltocephalus Dumeril & Bibron, i.e. Tschudi.

1843. Otolophus "Cuv." Fitzinger, Syst. Eept., p. 32. Type by original
designation: Bufo margaritifer Daudin =: Eana typhonic Lin-
naeus.

1843. Eurhina Fitzinger, Syst. Eept., p. 32. Type by original designa
tion: Bufo prohoscideus Wagler := Eana typhonia Linnaeus.

1843. Chilophryne Fitzinger, Syst. Eept., p. 32. Type by original desig-
nation: Bufo d'oriignyi Dumeril & Bibron.

1843. Phryne Fitzinger, Syst. Eept., p. 32. Type by original designation :
Bufo vulgaris Laurenti = Eana bufo Linnaeus.

1845. Anaxyrus Tschudi, Fauna Peruana, Herp., p. 78. Type by mono-
typy: A. melancholicus Tschudi = Bufo compactilis Wiegmann.

1849. Schismaderma A. Smith, Illus. Zool. S. Africa, Eept., App., p, 28.
Type by original designation : S. lateralis Smith = Bufo carens
Smith.

1860. Adenomus Cope, Proe. Acad. Nat. Sci. Philadelphia, p. 371. Type
by monotypy: A. badioflavus Cope = Bufo Tcelaarti Giinther.

1862. Elmebo Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 357. Type by
monotypy: Bufo haematiticus Cope.

1865. Epidalea Cope, Nat. Hist. Eeview, 5, p. 102. Type by monotypy:
Bufo calamita Laurenti.

1868. Otaspis Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 312. Type
by monotypy : Peltaphryne empusa Cope.

1868. Pegaeus Gistel, in Blicke, Leben Natur. Menschen (Leipzig), p.

310 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

161. Type by designation of Mertens:1936: Bona bufo Linnaeus.
1870. Nannophryne Giinther, Proc. Zool. Soc. London, p. 401. Type by

monotypy: N. variegata Giinther.
1870. Ansonia Stoliczka, Journ. Asiatic See. Bengal, 39, p. 152. Type by

monotypy: A. penangensis Stoliczka.
1876. Cranopsis Cope, Journ. Acad. Nat. Sci. Philadelphia, (N.S.) 8,

p. 96. Type by monotypy: C. fastidiosus Cope.
1876. Crfipidius Cope, Journ. Acad. Nat. Sci. Philadelphia, (N.S.) 8, p.

97. Type by monotypy: C. epioticus Cope.
1876. OUotis Cope, Journ. Acad. Nat. Sci. Philadelphia, (N.S.) 8. p. 98.

Type by monotypy: 0. caerulescens Cope.
1879. Dromoplectus Camerano, Atti. R. Acad. Sci. Torino, 14, p. 882.

Type by monotypy: Bufo anomalus Giinther = B. compactilis

Wiegmann.

Bufo cccrens Smith

1848. Bufo car.ens A. Smith, lUus. Zool. S. Africa, Rept., pi. Ixviii, fig,

1 : Interior of South Africa.

1849. Schismaderma lateralis A. Smith (new name for Bufo car ens

Smith), Illus. Zool. S. Africa, Rept., App., p. 28.
Range. Kenya Colony and Tanganyika Territory, south to
Natal, northwest through Bechuanaland to the Belgian Congo.

Bufo brauni Nieden Dead-leaf Toad

1910. Bafo braimi Nieden, Sitzb. Ges. Naturf. Freunde Berlin, p. 450:
Amani, Usambara Mountains, Tanganyika Territory.

Range. Mrgin forests of the Usambara ; Magrotto and Ulu-
guru Mountains, Tanganyika Territory.

Bufo regularis reguloris Reuss Egyptian Square-marked Toad
1834. Bufo regularis Reuss, Mus. Senckenberg., 1, p. 80: Egypt.
1841. Bufo pantherinus Dumeril & Bibron (part)^®^ Erpet. Gen. 8. j).

687: Egypt (A. Lefebvre coll.) by present restriction.
1855. Bufo mosaica Seetzen, Reise Syria, Palast., Aegypt. . . ., 3, p. 492:

Egypt.
1858. Bufo guineensis Giinther (part). Cat. Batr. Sal. Brit. Mus., p. 59:

Coast of Guinea (restricted).
1868. Bufo spinosus Bocage (not of Daudin:1803), Proc. Zool. Soc.

London, 1867, p. 845: Benguela, Angola.
1897. Bufo garmani Meek, Field Mus. Nat. Hist. Zool. Series, 1, p. 176:

* ' Haili, ' ' i.e. Halleh, British Somaliland.

187 Sometimes attributed to Boie, but his pantherinus, as well (is its use by
Tschudi (1838), constitute nomina ituda.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 311

1932. Bufo regularis kisoloensis Loveridge/®^ Occ. Papers Boston Soc.
Nat. Hist., 8, p. 52: Kisolo, Kigezi district, Uganda.

Range. Except in the northeast (Morocco and Tunisia), pres-
ent in all African countries from Algeria (Hoggar and A'ir)
and Libya (Fezzan) east to Egypt, south through Uganda, Kenya
Colony, Tanganyika Territory, Pemba and Zanzibar Islands,
to Natal and the Cape, though in some areas as more or less
recognizable races, of which many have been described.

Bufo regularis marakwetensis Roux^*''

1935. Bufo regularis marakivetensis Eoux, Mission Sci. de I'Omo, 3,

Zool., p. 180: Mount Marakwet at 2500 metres, Kenya Colony.
Range. Kenya Colony (known only from the two type S S ).

Bufo funereus Bocage

1866. Bufo funereus Bocage, Jorn. Sci. Lisboa, 1, p. 77: Duque de

Braganca, Angola.
1882. Bufo henguelensis Boulenger, Cat. Batr. Sal. Brit. Mus., pp. 299,

475, pi. xix, fig. 3: Benguela, Angola (restricted).
Range. Uganda, southwest to Angola ; northwest to Cameroon
and Fernando Po (also Dahomey according to Chabanaud :1916).

•

Bufo camenuiensis camerunensis Parker

1936. Bufo camerunensis camerunensis Parker, Proe. Zool. Soc. London,

p. 153 : Oban, Calabar, Nigeria
Range. Uganda, west to Liberia (but as yet unrecorded from
Dahomey and Togo).

Bufo latifrons Boulenger Spiny-flanked Toad

1900. Bufo latifrons Boulenger, Proc. Zool. Soc. London, p. 435, pi.
xxvii, fig. 1 : Benito River, Gabon, French Equatorial Africa.
Range. Kenya Colony;"" French Congo; French and British
Cameroons.

188 Revived by Laurent (1952, Herpetologica. 8, p. 53) as a full species sym-
patric with regularis. After careful reappraisal of the paratypes, together with
the large series that led me to synonyinize kisoloensis, I find Laurent's allegedly
diflerential characters offer no grounds for recognizing kisoloensis.

189 Without topotypical material it is unwise to suggest that this is a recogniz-
able race, a full species, or perhaps a synonym of regularis or some allied
species

190 The inclusion of this species rests on a toad taken on Mount Kinangop
between 2600 and 2700 metres, according to Angel (1925, Rept. et Batr., p. 56,
in "Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale (1911-1912).
Resultats Scientifiques. Vertebrata" (Paris). At my request Mons. Guib6 has
kii.dly re-examined this So (not 55) mm. specimen, and states (19.1.1957) that
in all respects it agrees with the deflnition of latifrons set forth in Parker's
(1936, Proc. Zool. Soc. London, p. 153) key, which should be consulted to avoid
confusion with B. c. camerunensis Parker.

312 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Bufo steindachnerii Pfeffer

1893. Bufo Steindachnerii Pfeffer, Jahrb. Hamburg. Wiss. Anst., 1892,
10, p. 103, pi. ii, fig. 8: Kihengo, Tanganyika Territory.

Range. Somalia, south through coastal Kenya Colony to Tan-
ganyika Territory.

Bufo vittatus Boulenger

1906. Bufo vittatus Boulenger, Proc. Zool. Soc. London, p. 573, fig. 98:
Entebbe, Uganda.

Range. Lower Egypt ; Uganda and western Tanganyika Ter-
ritory.

Bufo parkeri Loveridge

1932. Bufo parkeri Loveridge, Bull. Mus. Comp. Zool., 72. p. 382:
Mangasini, Usandawi, Tanganyika Territory.

Range. Tanganyika Territory (known only from the type
series).

Bufo urvmguensis Loveridge

1932. Bufo urunguensis Loveridge, Bull. Mus. Comp. Zool., 72, p. 383:
Kitungulu, Urungu, Tanganyika Territory.

Range. Tanganyika Territory (known only from the type
series).

Bufo ushoranus Loveridge

1932. Bufo ushoranus Loveridge, Occ. Papers Boston Soc. Nat. Hist., 8,
p. 45: Ulugu, Ushora, Tanganyika Territory.

Range. Tanganyika Territory (known only from the type
series).

Bufo lindneri Mertens

1955. Bufo lindneri Mertens, Jahrb. Ver. Vaterl. Naturk. Wurttemberg,

110, p. 48: Dar es Salaam, Tanganyika Territory.
Range. Tanganyika Territory (known only from the type S

and 9 ).

Bufo lonnbergi lonnbergi Andersson

1911. Bufo lonnhergi Andersson, Svenska Vetensk.-Akad. Handl., 48,

No. 8, p. 35, pi. ii, figs. 4a-b and 6a-b: Mount Kenya, Kenya

Colony.
Range. Kenya Colony.

Bufo lonnbergi nairobiensis Loveridge ^^^

191 On geographical grounds this race Is very doubtfully distinct, since it has
now been taken on Mount Kinangop and the Mau.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 313

1932. Bufo lonnbergi nairobiensis Loveridge, Occ. Papers Boston Soc.

Nat. Hist., 8, p. 48: Nairobi, Kenya Colony.
Range. Kenya Colony.

Bufo mocquardi Angel

1924. Bufo mocquardi Angel, Bull, Mus. Hist. Nat. Paris, 30, p. 269:

Mount Kenya and Mount Kinangop, Kenya Colony.
Range. Kenya Colony.^^"

Bufo taitonus taitanus Peters ^''^

1878. Bufo taitanus Peters, Monatsb. Akad. Wiss. Berlin, p. 208, pi. ii,
fig. 8: "Taita," i.e. Teita, Kenya Colony,

Range. Kenya Colony; Tanganyika Territory (Iramba Pla-
teau) ; Mozambique (Chifumbazi).

Bufo taitanus uzunguensis Loveridge

1932. Bufo taitanus uzunguensis Loveridge, Occ. Papers Boston Soc. Nat.
Hist., 8. p. 44: Kigogo, Uzungwe Mountains, Tanganyika Terri-
tory.
Range. Uzungwe, Ubena and Poroto Mountains, southern
Tanganyika Territory.

Bufo micranotis micranotis Loveridge

1925. Bufo micranotis Loveridge, Proe. Zool. Soc. London, p. 770, pi. i,

fig. 1 : Kilosa, Tanganyika Territory.
Range. (Possibly Coastal Kenya Colony on basis of tadpoles
from Diani Beach) ; Tanganyika Territory.

Bufo micranotis rondoensis Loveridge

1942. Bufo micranotis rondoensis Loveridge, Bull. Mus. Comp. Zool., 91.
p. 387: Nchingidi, Rondo Plateau, Tanganyika Territory.

Range. (Coastal Kenya Colony if Diani Beach tadpoles are
referable to this race) ; Hondo Plateau, Tanganyika Territory;
•langombe Swamp, Zanzibar (in M.C.Z.).

Bufo anotis Boulenger

li'07. Bufo anotis Boulenger, Ann. Mag. Nat. Hist., (7) 20. p. 48, pi.
iii: Chirinda Forest, southeast Mashonaland, Southern Rhodesia.

192 The Angola record of Menard (1937) is certainly erroneous. This little
toad is quite distinct from nairobiensis.

193 Many species or races of earless toads have been recorded under this name
from neighbouring territories, including Songosongo Island, Tanganyika Terri-
tory. All such require careful reappraisal before being included in the range.
Other races occur in Nyasaland and Mozambique.

314 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Range. Kilwa, Tanganyika Territory and Southern Rho-
desia.^^*

Genus NECTOPHRYNOIDES Noble

1926. Nectophrynoides Noble, Amer. Mus. Novit., No. 237, p., 15. Type
by original designation: Nectophryne tornieri Koux.

Nectophrynoides tornieri (Ronx) Usambara Viviparous-Toad
1906. Nectophryne tornieri Koux, Proc. Zool. Soc. London, 1, p. 63, pi.
ii, fig. 4: Ukami, i.e. Uluguru Mountains, Tanganyika Terri-
tory.
1910. Nectophryne wcrthi Nieden, Sitzb. Ges. Naturf. Freunde Berlin,

p. 439 : Dar es Salaam, Tanganyika Territory.^^^
Range. Usanil)ara ; Magrotto and Uluguru Mountains, Tan-
ganyika Territory.

Nectophrynoides vivipara (Tornier) Rungwe Viviparous-Toad

1905. Pseudophrync vivipara Tornier, Sitzb. Akad. Wiss. Berlin, 2, p.
855 : Rungwe and Kinga Mountains, Tanganyika Territory
( restricted ).-^^^
Range. Uluguru ; Uzungwe ; Ukinga ; Rungwe and Poroto
Mountains. Tanganyika Territory.

Suborder DIPLASIOCOELA
Family RHACOPHORIDAE'*
Genus CHIROMANTIS Peters

1855. Chiromantis Peters, Arch. Naturg., 21, Abt. 1, p. 56. Type by
monotypy: C. xerampelina Peters.

194 My early (1925) record of the occurrence of this species in western Tan-
ganyika Territory was based on misidentifled vittatus (Biikoba) or the subse-
quently described iishoraHus (Ushora).

195 Dar es Salaam is rejected pending confirmation. It may be observed that
botli wcrthi and some of the cotypes of viripara were allegedly collected in Dar
es Salaam by Werth. Werth was a chemist who had a shop in Dar es Salaam.
Possibly, like other residents at the coast, he may have spent a vacation at one
or other of the mission stations in the nearby Uluguru Mountains where both
species occur — and failed to label his specimens. Alternatively some of these
arboreal toads may liave been brought to Dar es Salaam in c;umps of bamboos
for planting in the Botanical Gardens. That they would survive is unlikely.

196 Includes POLYPEDATIDAE. Though included families are predominantly
diplasiocoelous. occasional memliers of the RHACOPIIORIDAE and MICROHY-
LIDAE have precocious vertebrae. Formerly the four African families of this
check list were grouped together as FIRMISTERNIA, but even that arrangement
had its arciferal exceptions.

Laurent (1951, Revue Zool. F>ot. Afr., 45, p. 120) suggests a new grouping In
which he would remove Hyperolius to a new family, the HYPEROLIIDAE, and
with it place such ranid genera as Astylosternus, Arthroleptis, Hemisus, etc.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 315

Chiromontis rufescens (Giiuther)
Western Foam-nest Tree-Frog
1868. Polypedates rufeseerm Giiuther, Proc. Zool. Soc. London, p. 486:

West Africa.
1875. Chiromantis guineensis Buchholz & Peters, Monatsb. Akad. Wiss.

Berlin, p. 203, pi. i, fig. 1 : Victoria, British Cameroon.
Range. Uganda,^^' west through the Belgian Congo to Nigeria.

Chiromantis xerampelina Peters

Southeastern Foam-nest Tree-Frog
1854. Chiromantis xerampelina Peters, Monatsb. Akad. Wiss. Berlin, p.

627 : Sena and Tete, Mozambique.
1920. Chiromantis umhelluzianus Ferreira, Joru. Sci. Lisboi (2), 8. p.
205, pis. i-ii: Umbelluzi Bridge, Lourengo Marques district,
Mozambique.
Range. Coastal Kenya Colony ; Tanganyika Territory and
Zanzibar Island, south through Mozambique, Nyasaland and the
Rhodesias to Zululand, Natal.

Chiromantis petersii kelleri Boettger
Northern Foam-nest Tree-Frog
1893. Chiromantis Tcelleri Boettger, Zool. Anz., 16, p. 131: Lake steppe
near Laku, and "Abdallah, " i.e. Abdulla country north of
Webi Valley, Ethiopia.
1900. Chiromantis hachowshii Nikolsky, Ann. Mus. Zool. Acad. Sci. St.

Petersbourg, 5, p. 246: Ferad, Ethiopia.
1916. Hylamhatcs cnantiodactylics Calabresi, Monit. Zool. Ital. (Fir-

enze), 27. p. 36, pi. ii, fig. 2: Bardera, Somalia.
1929. Chiromantis maerops Ahl, Zool. Anz., 80. p. 29: Ganda Ali,

"Annia" i.e. Ennia Galla, Ethiopia.
Range. Ethiopia and British Somaliland south through So-
malia to uorthern Kenj'a Colony.

Chiromantis petersii petersii Boulenger
Central Foam-nest Tree-Frog
1882. Chiromantis petersii Boulenger, Cat. Batr. Sal. Brit. Mus., p. 93,
pi. X, figs. 1-la: "Interior of East Africa," i.e. Mpwapwa,
Ugogo, Tanganyika Territory.

19' The only Uganda record is that of Loveridge (1942) for a nest taken in
Budongo Forest, northwestern Uganda. In 1902 Boulenger listed acerampcUna
from Uganda (apparently without supporting material), but it is far removed
from the range of that species.

316 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1929. Chiromantis albescens Ahl, Zool. Anz., 80, p. 30: Pokomoni, (oppo-
site Lamu Island), Kenya Colony.

1929. Chiromantis fasciatus Ahl, Zool. Anz., 80, p. 31 : Teita, Kenya

Colony; also Ikoma, Tanganyika Territory.

1930. Chiromantis pygm/ieus Ahl, Zool. Anz., 88, p. 219: Kibwezi, Kenya

Colony.

1931. Cliiromantis pictus Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p. 213:

Kilimatinde, Tanganyika Territory.
1931. Chiromantis rugosus Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p.

215: " Langenburg, " i.e. Manda, Lake Nyasa, Tanganyika

Territory.
Range. Kenya Colony (south of the equator), southwest to
central Tanganyika Territory .^^*

Genus LEPTOPELIS Giinther

1858. Leptopelis Giinther, Cat. Batr. Sal. Brit. Mus., p. 89. Type by

monotypy: Hyla aubryi A. Dumeril.
1924. Pseudocassina Ahl, Mitt. Zool. Mus. Berlin, 11, p. 8. Type by

monotypy: P. ocellata Ahl (not Hylambates ocellatus Mocquard)

as it includes P. rugosa Ahl {jide Ahl).
1941. Elaphromantis Laurent, Revue Zool. Bot. Afr., 35, p. 98. Type by

original designations: Hylambates notatus Peters.
1941. Ueteropclis Laurent, Revue Zool. Bot. Afr., 35, p. 99. Type by

original designation: Leptopelis parTceri Barbour & Loveridge.
1941. Taphriomnntis Laurent, Revue Zool. Bot. Afr., 35, p. 101. Type by

original designation: Cystignathus bocagii Giinther.
1950. Cryptothylax Laurent & Combaz, Revue Zool. Bot. Afr., 43, p. 276.

Type by monotypy: Hylambates greshoffl Schilthuis.

Leptopelis grcnnineus (Boulenger)^"''

1898. Megalixalus gramineus Boulenger, Ann. Mus. Civ. Stor. Nat.
Genova (2) 18, p. 721, pi. x, fig. 2: Between Badditu and
Dime, Somalia.

19!* It might be pointed out tliat Kenya is included solely on the basis of the
types of albescens, fasciatus and pygmaetis. Much of Ahl's material was sub-
adult and it was on the descriptions that I synonyuiized them with petcrsU in
19oo (Bull. Mus. Comp. Zool., 74, p. 390). When the opportunity occurs to exam-
ine the Pokomoni type it should be compared with C. p. killeri; also with xeram-
piiina which I obtained in series at nearby Witu. The Ikoma fasciatus is
certainly p. petersii, but the Teita specimen was apparently referred to xeram-
pclina by an earlier worker. Teita and nearby Kibwezi are faunistically similar,
so pygmaens is likely to be specifically identical with the Teita frog.

199 Admitted to this list on the somewhat slender evidence of four tadpoles
from Domorso waterhole near Moyale, recorded by Scortecci in 1940. In the same
paper he records four juveniles from Javello Plain near Agheremariani. The
only other Ethiopian record is that of Parker (1930) for Wourambouechi.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 317

Range. Ethiopia and Somalia, south to northern Kenya Col-
ony.

Leptopelis bocagii (Giinther)

1864. Cystignathus bocagii Giinther, Proe. Zool. Soc. London, p. 481, pi.

xxxiii, fig. 2: Duque de Braganca, Angola.
1893. Hylamhates angolensis Bocage, Jorn. Sci. Lisboa, (2) 3. p. 119:

Caeonda, Angola.
1893. Hylamhates cynnamomeus Bocage, Jorn. Sci. Lisboa, (2) 3. p. 120:

Quilengues, Angola.
1904. Hylamiates hocagei var. leiicopunctata Ferreira, Jorn. Sci. Lisboa,

(2) 7. p. 113: Gumba, Angola.
1937. Leptopelis hocagei haasi Mertens, Abhand. Senckenberg. Naturf.
Ges., No. 435, p. 21, fig. 2: Nsombo, Lake Bangweulu, Northern
Rhodesia.
Range. Kenya Colony (from Northern Uaso Nyiro), south
through Tanganyika Territory to Nyasaland, west through
Northern Ehodesia to Southwest Africa, north through Angola
to the Belgian Congo.

Leptopelis anchietae (Bocage)

1873. Eylambatcs anchietae Bocage, Jorn. Sci. Lisboa, 4, p. 226: Mos-

samedes, Angola.
1895. Hylamhates marginatus Bocage, Herp. Angola Congo, p. 178:

Quisange, Benguela, Angola.
1930. Hylamhates hrevipalmatus Ahl, Zool. Anz., 87, p. 228: Unyika,

Tanganyika Territory.
Range. Southwestern Tanganyika Territory ,^°° west to Angola,
north through Belgian Congo to the Bamenda District of French
Cameroon.

Leptopelis argenteus (Pfeffer)-"^

1893. Hylamhates argenteus Pfeffer, Jahrb. Hamburg. Wiss. Anst., 1892,
10, p. 100, pi. ii, fig. 3: Marsh south of Bagamoyo, Tanganyika
Territory.
Range. Coastal (Bagamoyo to Kuponda) Tanganyika Terri-
tory.

200 Known onlj- from a single record from the southwest (Ahl's type of brevi-
paltnatus) for the frogs assigned to anchietae by Tornier (1896) were made
paratypes of signijer by Ahl =; lermiculatus (Boiilenger).

201 Erroneously referred to the synonymy of Hylamhates maculatus by me
(1930, Proe. Zool. Soc. London, p. 24).

318 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Leptopelis concolor Alil"°"

1929. Leptopelis concolor Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p.
129: "Wito," i.e. Witu, Kenya Colony.

Range. Coastal Kenya Colony, south to Mikindani, Tan-
ganyika Territory.

Leptopelis flavomaculatus (Glintlier)^**'^

1864. Hyperolius flavomaculatus Giinther, Proc. Zool. Soc. London, p.

310, pi. xxvii, fig. 1: Eovuma Bay, Tanganyika Territory.
1897. Hylamhates johnstoni Boulenger, Proc. Zool. Soc. London, pp. 801,
803, pi. xlvi, fig. 4: Kondowe to Karonga, Nyasaland (re
stricted).
Range. Kenya Colony (Ngatana), south through Tanganyika
Territory to Mozambi(iue, west through Nyasaland to Southern
Rhodesia; also eastern Belgian Congo.

Leptopelis aubryi (Dumeril)

18.14. Hyla punctata Hallowell (not of Schneider : 1799), Proc. Acad.

Nat. Sci. Philadelphia, p. 193 : near Gabon River, French Congo.
1856. Hyla aubryi A. Dumeril, Revue Mag. Zool., (2) 8, p. 561: Gabon,

French Congo.
1929. Leptopelis harhouvi Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p.

119: Mount Lutindi, Usanibara Mountains, Tanganyika Terri

tory.2o*

Range. Usanibara Mountains, Tanganyika Territory ; Belgian
Ruanda-Urundi, west to Angola, north to Cameroon, west (with
gaps) to French Guinea.

Leptopelis notolus christyi (Boulenger)

1906. Uylamhates cuhito-alhiis Boulenger (part), Ann. Mag. Nat. Hist.,

(7) 17. p. 323: only the juvenile eotypes from Lake Albert,

Bunyoro, Uganda.-"''
1912. Hylamhates christyi Boulenger, Ann. Mag. Nat. Hist., (7) 12, p.

556 : Mabira Forest, ' ' Chagwe, ' ' i.e. Kyagwe, Uganda.
1929. Leptopelis budducnsis Ahl, Sitzb. Ges. Naturf. Freunde Berlin,

p. 119: Northwest Buddu Forest, "Deutsch-Ost-Afrika," i.e.

Uganda.

202 The Mombasa record of nattilensis (Smith) by Peters (1869) is probably
referable to concolor, the two species being superficially very alike.

203 Based on a juvenile, while the synonym was described from an adult ; it
serves to emphasize the strilving age variation presented by members of this
genus.

204 The types (M.C.Z. 13561-71), which neither I nor others who have examined
them can distinguisli from Cameroon aubryi except in size, were never seen by Ahl
who had no representatives of the "species."

205 Tile name cuhito-alhus should be restricted to the eotypes from Zima.
Cameroon

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 319

Range. Virgin forests of Uganda.

Leptopelis vermiculotus (Boiilenger)"""

1909. nylambatcs vcrmiculatus Boulenger, Ann. Mag. Nat. Hist., (8) 4,

p. 497: Amani, Usambara Mountains, Tanganyika Territory.

1929. Leptopelis sl(jnifer Ahl, Sitzb. Ges. Naturf. rreunde Berlin, p.

216: Devema, Usambara Mountains, Tanganyika Territory.
Range. Virgin forests of the Usambara, Magrotto and Rimgwe
Monntains, Tanganyika Territory.

Leptopelis karissimbensis Ahl

1929. Leptopelis Tcarissimhfinsis Ahl, Sitzb. Ges. Naturf. Freunde Berlin,

p. 195: Mtulia Gama, Mt. Karisimbi, Belgian Euanda.
1929. Leptopelis Icivuensis Ahl, Sitzb. Ges. Naturf. Freunde Berlin, i>.

206 : Kisenyi, Lake Kivu, Belgian Ruanda.
1929. Leptopelis graueri Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p. 213:

70 km. west of south end of Lake Edward, Belgian Congo.
1929. Leptopelis rugegensis Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p.

218: Rugege Forest, Belgian Ruanda.
Range. Virgin forests of southwest Uganda; Belgian Ruanda-
Trundi and the Belgian Congo.

Leptopelis parkeri Barbour & Loveridge ""'

1928. Leptopelis parJceri Barbour & Loveridge, Mem. Mus. Comp. Zool..

50, p. 236, pi. iv, figs. 9-10: Vituri, Uluguru Mountains, Tan-
ganyika Territory.

1929. Leptopelis martiensseni Ahl, Sitzb. Ges. Naturf. Freunde Berlin.

p. 219 : ? Amani or Tanga, Tanganyika Territory.
Range. Virgin forests of ITsambara and Uluguru Mountains,
Tanganyika Territory.

Leptopelis ulugxuxiensis Barbour & Loveridge

192S. Leptopelis ulugnrKeitsis Barbour & Loveridge, Mem. Mus. Comp.

Zool., 50, p. 235, pi. iii, fig. 3 : Nyange, Uluguru Mountains,

Tanganyika Territory.
1929. Leptopelis amaniensis Ahl, Sitzb. Ges. Naturf. Freunde Berlin,

p. 197 : Amani, Usambara Mountains, Tanganyika Territory.
1929. Leptopelis usamharae Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p.

205: Usambara Mountains, Tanganyika Territory.
1929. Leptopelis tanganus Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p.

221: Tanga, Tanganyika Territory.

206 The numerous specimens from U.sambara localities listed as rujus Reiche-
iiow by Nieden (191.5) are referable to vermiculatiis.

207 Some early and one recent record of cM/f/s Reichenow are referable to this
species.

320 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1929. Leptopelis grandiceps Ahl, Sitzb. Ges. Naturf. Freunde Berlin,

p. 207 : Amani, Usambara Mountains, Tanganyika Territory.
Range. Virgin forests of Usambara and Uluguru Mountains,
Tanganyika Territory.

Genus HYLAMBATES Dumeril

1858. Eylambates A. Dumeril, Ann. Sei. Nat., (3) 19. p. 162, Type by

monotypy: H. maculatus A. Dumeril.
1950. Phlyctimantis Laurent & Combaz, Revue Zool, Bot. Afr., 43. p,

274: Type by original designation: Eylambates leonardi Bou-

lenger.

Hylambotes maculatus Dumeril Red-blotched Black Frog
1853. Hylamhates maculatus A. Dumeril, Ann. Sci. Nat., (3) 19. p. 165,

pi. vii, figs, lib and 4: Zanzibar Island.
1944. Kassina poweri Hoffman, Soolog. Navors. Nas. Mus. Bloemfontein,

1, p. 169, figs. 1-3: St. Lucia, Estuary, Zululand.
Range. Kenya Colony ; Tanganyika Territory ; Zanzibar
Island ; Mozambique and Nyasaland ; Zululand, Natal.

Hylambates verrucosus Boulenger

1912. Ilylambates verrucosus Boulenger, Ann. Mag. Nat. Hist., (8) 10.
p. 141: Mabira Forest, "Chagwe, " i.e. Kyagwe, Uganda.

Range. Uganda and eastern Belgian Congo (fide No])le; also
Witte).

Genus KASSINA Girard

208

1853. Kassina Girard, Proc. Acad. Nat. Sci. Philadelphia, p. 421. Type
by monotypy: Cystignathus senegalensis Dumeril & Bibron.

1864. Cassina Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 182. Emenda-
tion for Kassina Girard.

1907. Paracassina Perraca, Boll. Mus. Zool. Anat. Comp. Univ. Torino,
22. No. 553, p. 3. Type by misidentification : Cystignathus sene-
galensis Dumeril & Bibron.^o^

1924. ITornierella Ahl, Mitt. Zool. Mus. Berlin, 11. p. 10. Type by subse-
quent designation of Ahl: 1926: T. pulclira Ahl = ?

208 This genus, so frequently redescribed on the basis of juveniles or adults in
which the vomerine teeth are absent, is much in need of a comprehensive
revision.

209 The description of Paracassina was based on two frogs (senegalensis)
from Toro, Uganda, which Peracca had misidentifled as Cassitia obsciira Boulen-
ger. Consequently, as pointed out by Pariser (1930, Proc. Zool. Sec. London, p.
31), Paracassina cannot be referred to the synonymy of Mocquardia Ahl (new
name for Rothschildia Mocquard, preoccupied by Rothschildia Grote:lS96).

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 321

1937. Cassiniopsis Monard, Bull. Soc. Neuchatel. Sci. Nat., 57. pp. 13,
41. Type by original designation: C. kuvangensis Monard.

1939. Scmnodactyhts Hoffman, Soolog. Xavors. Nas. Mus. Bloemfontein,

1, p. 89. Type by monotypy: S. thahanchuensis Hoffman.

1940. Kassinula Laurent, Revue Zool. Bot. Afr., 33, p. 313. Type by

original designation: K. ivittei Laurent = Cystignathus sene-
galensis Dumeril & Bibron (juv.).

Kassina senegalensis (Dunieril & Bibron )-'°

1841. Cystignathus ScJicgalensis Diuneril & Bibron, Erpet. Gen., 8, p.

418: Galam Lakes, Senegal.
1854. Cystignathus argyreivittis Peters, Monatsb. Akad. Wiss. Berlin,

p. 626 : Cabaceira, Mozambique.
1917. Megali.ralus macnlosu-s Sternfeld, Wiss. Ergebn. Zweiter Deutseh.

Zentral-Afrika-Exped. 1910-1911, 1, p. 501: Duma, Ubangi,

French Equatorial Africa.-'^
1930. Kassina descrticola Alil, Zool. Anz., 83. p. 280: Windhuk, South-
west Africa.
1930. Kassina modesta Ahl, Zool. Anz., 88. p. 281; Marianhill, Natal.
1933. Cassina angeli Witte, Revue Zool. Bot. Afr., 23. p. 172: Lukafu,

Kundelungu, Belgian Congo. '^-
1940. Kassinula ivittci Laurent, Revue Zool. Bot. Afr., 33. p. 314, figs.

lb and 2 (based on juveniles of from 12.5 to 14 mm.) : Kan-

senia, Belgian Congo.
1942. Kassina senegalensis ovambocnsis Hoffman, Soolog. Navors. Nas.

Mus. Bloemfontein, 1. pp. 150, 155, figs. 26-27: Ovamboland,

Southwest Africa.
Range. Sudan, east to Ethiopia (and possibly British Somali-
land), south through Uganda, Kenya Colony and Tanganyika
Territory to Natal, west to Cape Province, north to the Cain-
eroons, west (with some gaps) to Senegal.

Genus AFRIXALUS Laurent -^^

-10 Dr. V. FitzSimons informs me (22..X.56) that he regards Cassina s. inter
media Werner (1898) as a synon.vm of wcalii Boulenger : while A', poiceri Iloff
man (1944) was based ou a specimen of Ili/lamhatiS maculatus.

211 Dr. R. Mertens has found the type of maciilosus to be a juvenile senegalen-
sis.

212 Dr. K. Laurent recentl.v (1954) revived angeli (which I synonymized in
193(5) as a subspecies, but on grounds that appear imlikely to stand the test of
a thorough, continent-wide study of this group. Such a revisionary examination
may well result in additions to the foregoing synonymy.

213 Originally proposed as a subgenus without designation of type. Whether
there is suthcient justitication for the separation by Laurent of Acanthixalus
(for spinosus Buchholz & Peters) or Hctaixalas (for Malagasy members for-
merly assigned to Mcgali.valus) remains to be seen. Megalixalus is certainly
monotypic with seijchillensis as the only species.

322 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1944. Afrixalus Laurent, Eevue Zool. Bot. Afr., 38. p. 113. Type by
present designation: M.egalixalus fornasinii congicus Laurent
= EyperoUus dorsalis Peters.

Afrixalus fornasini fornasini (Bianconi)
Mozambique Bauana-Frog
1849.-^'* Euchnemis Fornasini Bianconi, Nuovi Ann. Sci. Nat., (2) 10.
1848, p. 107, pi. V, fig. 1; also 18o0, Spec. Zool. Mozamb., Eept.,
p. 23, pi. V, fig. 1: Mozambique.
1854. Hyperolius hivittatus Peters, Monatsb. Akad. Wiss. Berlin, p. 627:

Boror, Mozambique.
1862. Hyperolius spinifrons Cope, Proc. Acad. Nat. Sci. Philadelphia,

p. 342 : Umvoti, Natal.
1913. Megalixalus fornasinii var. unicolor Boettger, in Voeltzkow, Reise

in Ostaf rika, 3. p. 349 : Pemba Island, Zanzibar.
1920. Megalixalus loveridgii Procter, Proc. Zool. Soc. London, p. 418,

fig. 4 : Morogoro, Tanganyika Territory.
1930. Megalixalus dorsimaeulatus A hi, Sitzb. Ges. Naturf. Freunde

Berlin, p. 92: Magrotto, near Tanga, Tanganyika Territory.
Range. Coastal Kenya Colony ; Tanganyika Territory ; Pemba,
Zanzibar and Mafia Islands ; Mozambique ; Natal and extreme
eastern Cape Province (unless spinifrons proves to be a recogniz-
able race).

Afrixalus fornasini dorsalis ( Peters j Cameroon Banana-Frog

187.5. Hyperolius dorsalis (Schlegel) Peters, Monatsb. Akad. Wiss. Ber-
lin, p. 206, pi. i, fig. 2: Boutry, Ashanti, Gold Coast, and
Victoria, British Cameroon.

1889. Megalixalus schneideri Boettger, Ber, Senckenberg. Naturf. Ges.,
p. 276: Bonamandune (King Bell Dorf), Cameroon.

1941. Megalixalus fornasinii congicus Laurent, Revue Zool. Bot. Afr.,
35. p. 120, fig. 1: Buta, Uele, Belgian Congo.

1951. Afrixalus dorsalis regularis Laurent, Ann. Soc. Roy. Zool. Belgique,
82, p. 25, fig. 1 : Lambarene, Ogooue River, French Congo.

Range. Uganda, west through Belgian Congo to French
Guinea.

Afrixalus fulvovittatus leptosomus (Peters)^"

214 April, 1S40, according to Sherborn, but Bianconi (1S59, Mem. Accad. Sci.
Bologna, 10. p. 498) claims the name (with a single i ending) was first pub-
lished on 4.ii.l847 '

215 In earlier times there was much confusion regarding these frogs ; both
names were freely applied to specimens from the eastern seaboard which were,
in reality, A. b. brachi/cncmis Boulenger. The only places in East Africa where
I have encountered leptosomus is at Kaimosi. K. C., and Ujiji, T. T. The latter
were recorded by me as brachi/vnvmis, and I am indebted to Dr. Laurent for
examining them and pointing out that they are actually juvenile leptosomus.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 323

1877. Hyperolius Icptosomus Peters, Monatsb. Akad. Wiss. Berlin, p.

619, pi. — , fig. 5: Chinchoxo, Cabinda.
1931. Hr/pcroUus brevipalviatus Ahl, Mitt. Zool. Mus. Berlin, 17, p. 25:

Saiignielima, French Cameroon.
1941. MegaJixaUts leptosomus vpembae Laurent, Eevue Zool. Bot. Afr.,

35. p. 125, fig. 3: Nyonga, Lake Upemba, Belgian Congo.
Range. Sudan; Uganda; western Kenya Colony; western
Tanganyika Territory, west to Angola, north to the Cameroons,
and possibly Dahomey (fide Laurent). ^^^

Alrixalus brachycnemis brachycnemis (Boulenger)^^^
Short-limbed Banana-Frog
1896. Megalixalus brachycnemis Boulengcr, Ann. Mag. Nat. Hist., (6)

17, p. 403, pi. xvii, fig. 2: Chiradzulu, Nyasaland.
1931. Hyperolius pygmaeus Ahl, Mitt. Zool. Mus. Berlin, 17. p. 22:

Tanga, Tanganyika Territory.
1931. Eyperolius viultifasciatus Ahl, Mitt. Zool. Mus. Berlin, 17, p. 24:

Rungwe, Tanganyika Territory.
1931. Hyperolius ipianae Ahl, Mitt. Zool. Mus. Berlin, 17, p. 43: Ipiana,

Tanganyika Territory.
1931. Hyperolius unicolor Ahl, Mitt. Zool. Mus. Berlin, 17, p. 122:

Ipiana, Tanganyika Territory.
Range. Eastern Kenya Colony ; eastern Tanganyika Terri-
tory; Zanzibar and Mafia Islands; Mozambique; Nyasaland.

Afrixalus uluguruensis (Barbour & Loveridge)
Montane Banana-Frog
1928. Megalixalus uluguruensis Barbour & Loveridge, Mem. Mus. Comp.
Zool., 50, p. 231, pi. iii, fig. 2: Vituri, Uluguru Mountains,
Tanganyika Territory.
Range. Virgin forests of Usambara; Magrotto and Uluguru
Mountains, Tanganyika Territory.

216 A. /. fulvovittatue: (Cope), inc. rittiger Peters, ranges from northern
Nigeria west to Senegal.

«

217 Manv earlv East African records of '■fuliovittatus" and "leptosomus" refer
to this species. " Trinomials are used on account of the western A. b. wiedholzi
(Mertens :1938), of which Mcgalixaliis schoiitedenl Laurent is a synonym. As
the range of typical brachycnemis is largely conhned to the coastal region inland
to Lake Nvasa, I (195.3* was probably wrong in synonymizing the slightly
larger A. b. orophilus Laurent from the Central .\frican highlands.

324 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus HYPEROLIUS Rapp'^'

1842. Hyperolius Kapp, Arch. Naturg., 8. Abt. 1, p. 289. Type by mono-

typy : H. marmoratiis Rapp.
1864. Eappia Giinther, Zool. Kec, 1, p. 130. New name for Hyperolius

Rapp, thought to be preoccupied by Uperolia Gray.

A. Species represented in the Museum of Comparative Zoology
by material (often types, paratypes or topotypes) from those
countries indicated liy an asterisk (*) in Range.

Hyperolius viridiflavus viridiflavus (Dumeril & Bibron)

1841. Euon.emis viridiflavus Dumeril & Bibron, Erpet. Gen., 8, p. 528:
Ethiopia.

218 This, to me the most perplexing of all African genera, is still the least
understood and most unsatisfactory. In 1931 Hyperolius was reviewed by Ernst
Ahl, who described 98 additional species, many of which were composites of
distinct forms having somewhat similar patterns. .\hl failed to realize that
the juvenile livery of many "species" or races is indistinguishable, though dif-
fering niarivedly from that of their respective subadults, which again may be
totally unlike the mature frogs. The males of the latter sometimes have entirely
different markings from those of their own females.

In ]9;U Ahl published two voluminous papers, that in Das Tierreich (No.
5.'), Amphibia Anura, III, xvi + 477 pp.) bears the date "March 19.31," and,
having figures of the numerous species, is the more useful. The other (Mitt. Zool.
Mus. Berlin, 17, pp. 1-132), presumably intended to appear first for it carries
the abbreviation "spec, nov." after each of the new names, is dated "1 April
1931." From it one can obtain type localities and get an idea of what paratypic
material (in many instances mistakenly called "cotypen") was at the author's
disposal. The seemingly meticulous, and extremely detailed descriptions actually
do not cover the variations of Ahl's available material ; often they are flagrantly
careless and misleading. It is but fair to warn workers that the keys in Das
Tierreich are worse than useless, being based on characters of a trivial or variable
nature common to a host of forms that differ solely in their striking color
patterns. When Ahl's series of "cotypen" was adequate, a selection of them was
obtained in 1932 by exchange for the Museum of Comparative Zoology. The avail-
ability of this material made it possible for me to synonymize almost 50 of Ahl's
alleged species ; Laurent has similarly dealt with some others. We have not
always agreed : in some instances, no doubt, because of the composite nature of
Ahl's "species," its faded condition, or other cogent reasons.

Laurent, who has done more work on this genus than anyone, has himself
proposed at least GO new Hyperolius. Following a decade of bewildering nomen-
clatorial changes in which species were synonymized or revived as racial entities
of this or that form, which in turn was liable to be demoted to subspecific rank
of something else, Laurent (1951) endeavored to bring order out of chaos by
assigning many of these names to subspecific status of much earlier forms like
marmordtus, viridiflarus, etc. In preparing this check list I hoped to follow
Laurent in this laudable effort, but encountered so many geographical difficulties
that — in the interests of stability — it seemed advisable to adhere to binomials
except where I have previously used trinomials.

Departing from the format used elsewhere in this check list, I have divided
these sedge- or tree-frogs into two groups. The first (A) contains those species
or subspecies represented by material in the Museum of Comparative Zoology
(mdicated by an asterisk*) that, as of today, do not seem capable of further
synonymization. The ranges given have usually been restricted to the country
of origin of the type series and its synonyms, together with those countries
from which I have, or have seen, material whose identifications I have been able
to verify. From what I have said, I hope it will be clear to enthusiastic local
amateurs that ranges should not be extended by the inclusion of early extra-
territorial records of the same "species," even though identified by a competent
herpetologist of the period. Such book work merely perpetuates confusion.

To the second group (B, see p. 334) are assigned those "species," usually
known only from a single holotyi>e, that I have not seen. In some instances it is
possible that the species in question may not even be referable to Hyperolius,
whose horizontal pupil is the only external character differentiating it from
Afrixalua — whose pupil is vertical.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 325

Range. Ethiopia, south to Bannissa, Northern Frontier Prov-
ince, *Kenya Colony.

Hyperolius pachydenna "Werner ^^^

1908. Bappia pachyderma Werner, Sitzb. Akad. Wiss. Wien, 1907, 116,

Abt. 1, p. 1403: Goudokoro, Sudan.
1921. Rappia burgeoni Witte, R^vue Zool. Afr., 9, p. 19, pi. v, figs. 2-2b:

' ' Madyu, Uele ' ' = Gangula Madi on Uele River, midway

between Bambili and Niangara, Belgian Congo.
1925. Rappia Bossii Calabresi, Atti. Soc. Ital. Sci. Nat. (Milano), 64,

p. 121, fig. — : Upper Uele region, Belgian Congo.
1931. Hi/peroUus phrynoderma Ahl, Das Tierreich, no. 55, p. 344, fig.

218: No locality, but collected by Deutsche Zentral-Afrika

Expedition.
1931. Hyperolius ocidatus Ahl, Das Tierreich, no. 55, p. 377, fig. 252:

Balaibo, banks of Duki River, Belgian Congo.
Range. *Sudan; *Uganda; *Kenya Colony; *Belgian Cono;o.

Hyperolius picturatus Peters""

1875. Hyperolius picturatus "Schlegel" Peters, Monatsb. Akad. Wiss.

Berlin, p. 206, pi. ii, fig. 2: Boutry, Ashanti, Gold Coast.
1875. Hyperolitis nitidulus Peters, Monatsb. Akad. Wiss. Berlin, p. 209,

pi. iii, fig. 4: Yoruba, Lagos, Nigeria.
Range. *Western Kenya Colony ; *Uganda ; *Belgian Congo ;
*French Congo; Sao Thome; *French Cameroon; Togoland;
Gold Coast ; *Ivory Coast ! *Liberia ; *Sierra Leone.

Hyperolius marginatus Peters

1854. Hyperolius marginatus Peters, Monatsb. Akad. Wiss. Berlin, p.

627: Maqanga, Mozambique.
1931. Hyperolius pictus Ahl, Das Tierreich, no. 55, p. 301, fig. 176:

Crater Lake, Ngosi Volcano, Tanganyika Territory.
1931. Hyperolius ngoriensis Ahl, Das Tierreich, no. 55, p. 324, fig. 198:

Crater Lake, Ngosi Volcano, Tanganyika Territory,
Range. Montane marshes of southern * Tanganyika Terri-
tory ; Mozambique and Nyika Plateau, northern *Nyasaland.

Hyperolius goetzei Ahl

1931. Hyperolius goetaei Ahl, Das Tierreich, no. 55, p. 413, fig. 286:
Uhehe, Tanganyika Territory.

219 With respect to the first two names (Rappia pachyderma was based on a
juvenile of generalized appearance), I have followed Laurent (1951) who regards
pachyderma as a race of viridiflavus. Hitherto I have referred to this frog as
rosgii.

220 The alleged example from Pemba Island, reported by Boettger (1913), on
being re-examined by Mertens (1940) was found to be a juvenile Afrixalus f.
fornasini.

326 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Range. Uplands of *Kenya Colony and *Tanganyika Terri-
tory.

Hyperolius discodactylus Ahl

1931. Hyperolius discodactylus Ahl, Das Tierreich, no. 55, p. 363, fig.

239 : Rugege Forest, Belgian Euanda-Urundi, and west of Lake

Edward, Belgian Congo.
1931. Hyperolius alticola Ahl, Das Tierreich, no. 55, p. 379, fig. 255:

at 1800 metres, Euwenzori Mountains, Belgian Congo.
Rcmge. *Ruwenzori Mountains, Uganda; *Belgian Congo.

Hyperolius montanus (Angel)

1924. Rappia montana Angel, Bull. Mus. Hist. Nat. (Paris), 30. p. 269:
Mount Kinangop, Aberdare Mountains, Kenya Colony.

Range. Montane marshes of Mts. Kinangop and Kenya,
*Kenya Colony.

Hyperolius pantherinus (Steindachner)

1891. Megalixalus pantherinus Steindachner, Anz. Akad. Wiss. Wien, 28.
p. Ii2: Laikipia, Kenya Colony.

Range. Kenya Colony (known only from the holotype : photo
inM.C.Z.V

Hyperolius bayoni (Boulenger)"^

1911. Uappla bayoni Boulcnger, Ann. Mus. Civ. Stor. Nat. Genova, (3)

5. p. 1G8: Bulolo; Busu ; Enteblje; Jinja; Kabulamuliro ; and

Mbale, Uganda.
1931. Hyperolius stuhmarini Ahl, Das Tierreich, no. 55, p. 395, fig. 271:

Vitchumbi, south end of Lake Edward, Belgian Congo.
1940. Hyperolius ornatus Laurent, Eevue Zool. Bot. Afr., 34, p. 4:

Eutshuru, Belgian Congo.
Range. *Uganda, southwest to Belgian Ruanda-Urundi (also
reported from Mozambique by Parker in 1931).

Hyperolius kivuensis Ahl""'

1931. Hyperolius kivuensis Ahl, Das Tierreich, no. 55, p. 280, fig. 151
(34 mm. $, Kandt coll.): Lake Kivu, Belgian Congo.

1931. Hyperolius kwidjwiensis Ahl, Das Tierreich, no. 55, p. 296, fig. 172
(32 mm. S, Schubotz coll.): "Kwidjwi," i.e. Idjwi Island,

221 A 9 frog (M.C.Z. 10287) from the Uganda shore of Lake Eflward differs
slightly from a topotype stuhlmanni (M.C.Z. 2S126).

222 In 1951 Laurent regarded kicidjiciensis (with which he would include
kandt i) as distinct. He would transfer koehi to the synonymy of schubotzi Ahl
(in whose synonymy I place the 37 mm. macrodaeti/lus) . The series of frogs
from Nyamliolo, Lake Tanganyika, N. Rhodesia which in 19:50 I referred to
rhodoscelis in error, appear to be indistinguishable from Kivu frogs.

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS 327

Lake Kivii, Belgian Congo.
1931. Hyperolius kandti Ahl, Das Tierreich, no. 55, p. 327 (33 mm. S,

Kandt coll.) : Lake Kivu, Belgian Congo.
1931. Hyperolius l-oehli Ahl, Das Tierreich, no. 55, p. 405 (33 mm. S ,

Koehl coll.) : Kisenyi, Lake Kivu, Belgian Euanda-Urundi.
Range. *Soiitlnvest Uganda ; *western Tanganyika Territory ;
^Northern Rhodesia ; *eastern Belgian Congo ; *Belgian Ruanda-
Urundi.

Hyperolius variabilis Ahl""^

1931. Hyperolius variabilis Ahl, Das Tierreich, no. 55, p. 297, fig. 173:

restricted to Bukoba, Tanganyika Territory.
1931. Hyperolixis wettsteini Ahl, Das Tierreich, no. 55, p. 343, fig. 217:

Bukoba, Tanganyika Territory.
1931. Hyperolius mohasicus Ahl, Das Tierreich, no. 55, p. 360, fig. 236:

Lake Mohasi, Belgian Euanda-Urundi.
1931. Hyperolius monticola Ahl, Das Tierreich, no. 55, p. 376, fig. 251 :

"Niansa," i.e. Nyanza, Belgian Euanda-Urundi.
1931. Hyperolius irregularis Ahl, Das Tierreich, no. 55, p. 396, fig. 272:

Lake Mohasi, Belgian Euanda-Urundi.
Range. Montane meadows of southwest *Uganda; Belgian
Ruanda-Urundi and adjacent *Belgian Congo.

Hyperolius latifrons Ahl"-*

1931. Hyperolius latifrons Ahl, Das Tierreich, no. 55, p. 335, fig. 208:
Bamboo forest at 2400 metres, near Mhi-abu Clahama village.
Mount Karisimbi, Belgian Euanda-Urundi.
1931. Hyperolius Icarissimhiensis Ahl, Das Tierreich, no. 55, p. 348, fig.
223 : Bamboo forest at 2400 metres, near Mhcabu Gahama
village, Mount Karisimbi, Belgian Euanda-Urundi.
Range. *Western Tanganyika Territory ; Belgian Ruanda-
Urnndi and *Belgian Congo.

Hyperolius multicolor Ahl"-'

1931. Hyperolius multicolor Ahl, Das Tierreich, no. 55, p. 367, fig. 243:
Bamboo forest at 2400 metres, near Mhcabu Gahama village,

223 Considered a race of viridiflavus by Laurent (1951) who would add punc-
tntis-simus Ahl and flaroguttatus (part) to the above synonymy. While Ahl's
material was often composite I scarcely think it warrants such action ; however,
icettsteini and monticola are added fide Laurent.

224 Both latifrons and karissimbiensia were based on juveniles of 21 and 23 mm.
respectively. Laurent (1951) regards karissimbiensis as a distinct race of viridi-
flavus to which he would add multicolor Ahl.

225 Our cotype and series of multicolor from Lake Bunyoni (with whose identi-
tication Laurent concurs) do not support its synonymization with "karissimbien-
sis."

828 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Mount Karisimbi, Belgian Ruanda-Urundi.
Range. Montane meadows of southwest *Uganda; *Belgian
Ruanda-Urundi and adjacent Belgian Congo.

Hyperolius bitaeniatus Ahl"^''

1931. Eyperolhts hitaeniatvs Ahl, Das Tierreich, no. 55, p. 322, fig, 196:
Ukonde-Unyika, Tanganyika Territory.

Rcmge. Southwestern Tanganyika Territorj^ and nearby
*Northern Rhodesia.

Hyperolius fuelleborni Ahl"^

1931. Hyperolius fuellchorni Ahl, Das Tierreich, no. 55, p. 349, fig. 224:
' ' Langenburg, ' ' i.e. Manda, Lake Nyasa, Tanganyika Territory.

Range. Region around northern end of Lake Nyasa, southwest
*Tanganyika Territory.

Hyperolius mariae Barbour & Loveridge^^^

1928. Hyperolius mariae Barbour & Loveridge Mem. Mus. Comp. Zool.,
50, p. 217, pi. iii, fig. 1: Derenia, Usambara Mountains, Tan-
ganyika Territory.
1931. Hyperolius melanophthalmus Ahl, Das Tierreich, no. 55, p. 341,

fig. 215 : Zanzibar.
1931. Hyperolius renschi Ahl, Das Tierreich, no. 55, p. 397: Zanzibar.
Range. Coastal *Tanganyika Territory and *Zanzibar
Island."^

Hyperolius flavoguttatus Ahl'^°

1931. Hyperolius flavoguttatus Ahl, Das Tierreich, no. 55, p. 369, fig.

245 : Bukoba, Tanganyika Territory.
Range. *Kenya Colony and *Tanganyika Territory.

226 At one time (1943) Laurent thought bitaeniatus might be a synonym of
rhodoscelis Bonlenger (with which he correctly synonymlzes iinduJatus Boulen-
gcr). A comparison of our andidatus cotype with our cotype of hitaeniatiis con-
vinces me thpy are distinct. Wo have also a good series of quite typical
bitaeniatus from Abercorn. More recently (19.51) Laurent has, mistakenly I
think, referred bitaeniatus to the synonymy of H. mariae Barbour & Loveridge.

227 Laurent (1951) refers fuelleborni to the synonymy of nyassae Ahl. We have
cotypes of both and I cannot agree ; to my thinking "nyassae" is a synonym of
rhodoscelis (Boulenger).

228 Laurent (1951) adds noblel Ahl to the synonymy; I adhere to my (1936)
alignment of it with puncticulatus (Pfeffer).

229 My earlier record from Lake Nyasa was based on a mlsidentified fuelleborni.

230 AH that the Museum of Comparative Zoology has is a bleached cotype from
Bukoba ; a correctly colored topotype from Mount Kenya, and a frog from
Mount Mbololo somewhat doubtfully attributable to flavoguttatus. These do not
suggest that Laurent is correct in synonymizing flavoguttatus with variabilis Ahl.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 329

Hyperolius udjidjiensis AhP^^

1931. Hyperolms udjidjiensis Ahl (part), Das Tierreich, no. 55, p. 370,

fig. 246: restricted to Ujiji, Tanganyika Territory.
Range. * Tanganyika Territory.

Hyperolius striolatus Peters ^^^

1882. Hyperolius striolatus Peters, Sitzb. Ges. Naturf. Freunde Berlin,

p. 9: "Taita," i.e. Teita, Kenya Colony.
1902. Eappia ferniquci Mocquard, Bull. Mus. Hist. Nat. (Paris), 8, p.

407: Athi River, Kenya Colony.
1931. Hyperolius caeruleopunctatus Ahl, Das Tierreich, no. 55, p. 350,

fig. 225: Nairobi and Kibwezi, Kenya Colony.
1931. Hyperolius pulchromarmoratus Ahl, Das Tierreich, no. 55, p. 367,

fig. 242: "British East Africa" (Hiibner of Kibwezi).
1931. Hyperolius udjidjiensis Ahl (part). Das Tierreich, no. 55, p. 370,

fig. 246 (only the paratypes from Kibwezi, Kenya Colony).
1931. Hyperolius scheffleri Ahl, Das Tierreich, no. 55, p. 387: Kibwezi.

Kenya Colony.
Range. *Kenya Colony.

Hyperolius rhodoscelis (Boulenger)

1901. Eappia rhodoscelis Boulenger, Ann. Mus. Congo, (1) 2, fasc. 1,
p. 3, pi. ii, figs. 1-la: Pweto, Lake Mweru, Belgian Congo.

1901. llappia undulata Boulenger, Ann. Mus. Congo, (1)2, fasc. 1, p. 4,

pi. ii, fig. 2: Pweto and Lofoi, Lake Mweru, Belgian Congo.

1902. Eappia symetrica Mocquard, Bull. Mus. Hist. Nat. (Paris) 8. p.

408: Athi Eiver, Kenya Colony.

1920. Eappia platyrhinus Procter, Proc. Zool. Soc. London, p. 416, fig. 3 :
Nairobi, Kenya Colony.

1931. Hyperolius asper Ahl, Das Tierreich, no. 55, p. 310: Nairobi,
Kenya Colony.

1931. ^Hyperolius hreviceps Ahl, Das Tierreich, no. 55, p. 316, fig. 190
(at least the paratypes from Eldama Ravine, Kenya Colony,
are rhodoscelis): "Tschimbo" = Chimbo, Mozambique.

1931. Hyperolius nyassae Ahl, Das Tierreich, no. 55, p. 339, fig. 213:
"Langenburg, " i.e. Mauda, Lake Nyasa, Tanganyika Terri-
tory.

Range. ? *Southeru Sudan; *Kenya Colony; *Tanganyika
Territory; ? Mozambique; ? *Angola; southern *Belgian Congo.

231 From Ujiji we have a topotype series consisting of a ^ and five 9 9
that I collected In 1939. I regard them as distinct from striolatus with which
species Laurent (1951) would synonymize udjidjiensis (part). It seems ques-
tionable whether Ahl's cotype from Kibwezi is conspecific.

232 To the synonymy of striolatus, Laurent would add symetrica, platyrhinus,
asper and the Baringo specimen of breviceps Ahl. Geographically this seems
plausible but is incorrect unless the species is dichromatic.

330 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Hyperolius punctatissimus Ahl"^'^

]P31. Hyperolius punctatiftsiniun Alil, Das Tierreich, no. 55, p. 299, fig.

174: restricted to Bukoba, Tanganyika Territory.
Range. * Tanganyika Territory.

Hyperolius vermicularis Ahl""*

1893. Fappin vermiculata Pfeffer (not of Peters: 1882), Jahrb. Ham-
burg. Wiss. Anst., 1892, 10. p. 98, pi. i, fig. 12: Zanzibar.

1931. Hyperolius vermiailaris Ahl (new name for vermiculata Pfeffer,
preoccupied by an Angolan species), Das Tierreich, no. 55, p.
275, fig. 145.

Range. *Kenya Colony; Zanzibar Island.

Hyperolius marmoratus onunatostictus Laurent "'^^

1951. Hyperolius marmoratus ommatostictus Laurent, Ann. Soc. Roy.
Zool. Bclgique, 82. p. 36: "Kibouoto," i.e. Kibongoto, Mount
Kilimanjaro, Tanganyika Territory.
Range. ^Tanganjnka Territory.

Hyperolius orgentovittis Ahl"^''

1931. Hyperolius argcntovittis Ahl, Da.s Tierreich, no. 55, p. 345, fig.
220: Ujiji, Tanganyika Territory.

1931. Hyperolius callichromus Ahl (part), Das Tierreich, no. 55, p. 372,
fig. 248: Western banks of "Russisi, " i.e. Ruzizi River and
northwest shores of Lake Tanganyika, Belgian Congo.

233 Laurent (1951) would synonymize this also with i-ariabUis Ahl, but our
solitary specimen from Kahare. r.ukol)a District, seems to show relationship
with strioldtiis rather than with variahUis.

-31 Laurent (1931) would add sci-iptux Ahl lo the synonymy. However, I failed
to meet with vermicularis in the vicinity of Tanga (from whence came the holo-
type of scriptus), obtaining our abundant material from seven localities north
of Mombasa.

235 The Museum of Comparatve Zoology topotypes (M.C.Z. 21009-11) were
taken at the same time as the type series. Kibougoto is not very high, viz. 1300
metres, roughly 4223 feet.

-36 In their tibial pattern our long series of argcntorittis topotypes form a linU
with the speckled tibial pattern exhibited by our callicluomus cotypes (M.C.Z.
17630-1). In the opposite direction — on the southeast shore of I>ake Tanganyika
— the even larger series I captured at Nyamkolo are remarkably uniform, dis-
playing a single red-brown streak on the tibia that is only occasionally broken
up into two or three elongate blotches. Some of our cotypes of brieni Laurent
(1943, Ann. Mus. Congo Beige, (1) 4, fasc. 2, p. 119, fig. 30 only), viz ^ and 5

(M.C.Z. 26264, 26266) have a dorsal pattern, and differ but slightly in tibial
pattern, from argentovittis. Yet our other cotypes of brieni ^ and J (M.C.Z.

26263, 26205), also from Nyonga, Upper Luapula, have the vermiculated dorsal
patterns so admirably portrayed in Laurent's figure 37. Thej' are unquestionably
referable to the same form as the frogs from Kabengere. Belgian Congo, that I
have referred to graucri Ahl. Either brieni is a cuiiiposite. being part argcnto-
vittis and part graucri, or the species is dichromatic, in which case Laurent is
correct in adding graueri to the synonymy of ai gcntoritlis and brieni should gc
too. Our cotype (M.C.Z 17633) of decipiens Ahl from the Ruzizi lends no su]'
port to Laurent's synonymizing of that species with nrgcnturil tis.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 331

Range. *Tanganyika Territory; *Nortlieru Rhodesia; *Bel-
gian Congo.

Hyperolius puncticulatus substriatus Ahl

Northern Broad-striped Sedge-Frog
1931. Hypcroliios substriatus Ahl, Das Tierreich, no. 55, p. 358, fig. 234:

Magrotto Mountain, Tanganyika Territory.
Range. Coastal *Kenya Colony (Malindi), south through the
Usambara Mountains to Morogoro, *Tanganyika Territory ; *Zan-
zibar Island.'^'

Hyperolius puncticulatus puncticulatus (Pfeffer)^^®
Southern Broad-striped Sedge-Frog
1893. L'ap2)ia puricticulata Pfeffer, Jahrb. Hamburg. Wiss. Anst., 1892,

10. p. 31, pi. ii, fig. 2: Zanzibar.
1931. Hyperolius noblei Ahl, Das Tierreich, no. 55, p. 400, fig. 275:

Kihva, Tanganyika Territory.
Range. Zanzibar Island ; Uluguru and Rungwe Mountains of
*Tanganyika Territory ; Misuku ; Nyika and Mlanje Mountains
of *Nyasaland.

Hyperolius argus ahli Loveridge"^^

Northern Argus-spotted Sedge Frog
1936. Hyperolius ahli Loveridge, Bull. Mus. Comp. Zool., 79, p. 402 : Lake

Pcccatoni, northeast of Witu, Kenya Colony.
Range. Coastal *Kenya Colony, south to Lindi. *Tanganyika
Territory.

Hyperolius argus argus Peters"*"

Southern Argus-spotted Sedge-Frog
1854. Hyperolius arfjus Peters, Monatsb. Akad. Wiss. Berlin, p. 628
(l)ased on 9 > : Boror, Mozambique.

23T M.C.Z. 17162 from Mwera, Zanzibar Island, is undoubtedly referable to this
form. It remains to be seen which race is dominant on the island.

-38 I disagree with Laurent's action in removing noblei from the synonyiny of
puncticulatus to that of mariae Barbour & Loveridge. Uganda and other records
outside the range given here, are considered questionable.

239 For a coloured plate showing the striking sexual dichromatism of this raci'.
whose ^ is git't'u tlt'i'Ucd with tilacK. see l.oviTidg.- 1 1942. Hull. Mus. t'oinii
Zool., 91, pi. iii).

240 Admission to this list depends on the identification of the juveniles from
Mafia Island reported by Parker (1937), and from the J from Zanzibar recorded

by Laurent (1943) : both are somewhat questionable. My Tanganyika material
identified as nrqus by Procter (1920), and consequentlv some of my own, were
actually referable to p. substriatus Ahl (1931). See Parker (1931, Proc. Zool.
Soc. London, pi. i) for astouishiug age and sexual dichromatism in a near topo-
typic series.

332 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1854. Hyperolius fiavoviridis Peters, Monatsb. Akad. Wiss. Berlin, p.

628 (based on S ) : Boror, Mozambique.
1854. Hyperolius Tettensis Peters, Monatsb. Akad. Wiss. Berlin, p. 628

(based on $): Tete, Mozambique.
1893. Bappia platycephala Pfeffer, Jahrb. Hamburg. Wiss. Anst., 1892,

10, p. 96, pi. ii, fig. 2 (based on $) : Quilimane, Mozambique.
Range. *Mozambique ; *Nyasalancl.

Hyperolius concolor sonsibaricus (Pfeffer)
Northeastern Straw-or-Greeu Sedge-Frog
1893. Eappia sansibarica Pfeffer, Jahrb. Hamburg. Wiss. Anst., 1892.

10. p. 97, pi. ii, fig. 4: Zanzibar.
1931. Hyperolius ruhripes Ahl, Das Tierreich, no. 55, p. 361: Kililana,

mainland opposite Lamu Island, Kenya Colony.
Range. *Keiiya Colony ; ^Tanganyika Territory ; Zanzibar
Island.-"

Hyperolius concolor tuberilingvds Smith "*^
Southeastern Straw-or-Green Sedge-Frog
] 849. II yperolius tub.erilinguis A. Smith, Illus. Zool. S. Africa, Rept.,

App., p. 26: country eastward of Cape Colony, i.e. Natal.
1862. nyperolius coccotis Cope, Proe. Acad. Nat. Sci. Philadelphia, p.

342: Umvoti, Natal.
1864. Hyperolius citrinus Giinther (part), Proe. Zool. Soc. London, p.

311, pi. xxvii, fig. 2: restricted to "Zambezi Expedition."
1S67. ''.Hyperolius granulosus Peters, Monatsb. Akad. Wiss. Berlin, p. 891,

footnote: Mozambique.
1947. Hyperolius Jcivucnsis smaragdinxis Laurent, Ann. Mag. Nat. Hist.,

(11) 14. p. 292: Charre, Mozambique.
1947. Hyperolius sansiharicus lovcridgei Laurent, Ann. Mag. Nat. Hist..

(11) 14. p. 294: Kitaya, Rovuma River, Tanganyika Territory.
Range. Southern *Tanganyika Territory; Mozambique; *Ny-
asaland; *Natal.

Hyperolius porkeri porkeri Loveridge

Northeastern Brown-or-Green Sedge-Frog
1933. Hyperolius parTceri Loveridge, Bull. Mus. Comp. Zool., 74. p. 410:
Bagamoyo, Tanganyika Territory.

-11 The Uganda record of Roux (1910) must be considered doubtful.

242 The Charre series on which Laurent based his smaragdinus, were originallj
referred by Parker (1931) to concolor (Hallowell) of Liberia. With this deter-
mination I can And no fault as Liberian and Natal frogs appear Indistinguishable
though it appears advisable to treat them as distinct until we linow more of the
intermediate races — of which kivuensis may well be one. The 133 frogs from
Kitaya, on some of which Laurent based his loveridgei, were originally assigned
by me to c. oitrimis Giinther.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 333

Range. Coastal *Kenya Colony, south to Dar es Salaam,
*Tanganyika Territory.

Hyperolius porkeri rovumae Loveridge"^^

Southeastern Bro\vn-or-Green Sedge-Frog
1942. Hyperolius parTceri rovumae Loveridge, Bull. Mus. Comp. Zool.,
91. p. 410, pi. iii, figs. 5-6: Kitaya, Rovuma River, Tanganyika
Territory.
Range. Coastal (Kilwa to Kitaya) *Tanganyika Territory.

Hyperolius nasutus Glinther

Sharp-and-blunt -snouted Sedge-Frog
1864. Hyperolius nasutus Giinther, Proc. Zool. Soc. London, p. 482, pi.

xxxiii, fig. 3 : Duque de Braganca, Angola.
1895. Eappia pwnctulata Bocage, Herp. Angola Congo, p. 168: Banks

of the ' ' Quanza ' ' i.e. Cuanza River, Angola.
1901. Eappia grannlata Boulenger, Ann. Mus. Congo, (1) 2, fasc. 1, p. 4,

pi. ii, fig. 3 : Pweto, Lake Mweru, Belgian Congo.
1901. Eappia oxyrhynchus Boulenger, Ann. Mus. Congo, (1) 2, fasc.
1, p. 0, pi. ii, fig. 4: Pweto and Lofoi, Lake Mwenu, Belgian
Congo.
1931. ^Hyperolius petersi Ahl, Das Tierreich, no. 55, p. 274, fig. 144:

Mombasa, Kenya Colony.
1931. Hyperolius acuticeps Ahl, Das Tierreich, no. 55, p. 282, fig. 153:

TJkonde-LTnyika, Tanganyika Territory.
Range. Ethiopia {fide Parker) ; Uganda {fide Loveridge) ;
*Keuya Colony; ^Tanganyika Territory; *Nyasaland; *North-
ern Rhodesia; ^Southern Rhodesia; *Angola; *Belgian Congo;
French Cameroon {fide Mertens :1940) ; Liberia (as nasutus
subsp. Laurent :1951) ; French Guinea (as oxyrJujnchus subsp.
Laurent :1951).

Hyperolius pusillus (Cope)"*"* Transparent Pigmy Sedge-Frog
1862. Crumenifcra pusilla Cope, Proc. Acad. Nat. Sci. Philadelphia, p.

343 : Umvoti, Natal.
1864. Hyperolius microps Giinther, Proc. Zool. Soc. London, p. 311, pi.

xxvii, fig. 3 : Rovuma Bay, Tanganyika Territory.

243 The brown ^ and green J with white side-stripe are shown in color on

plate iii, together with the two sexes of the typical form which lacks a light side
stripe.

244 The heavily spotted milnei from northeast Kenya may appear to differ from
pusillus of Natal, but so many intermediate conditions occur along the 2000 miles
that separate them it seems advisable to ignore subspecies. Many West African
frogs have been referred to pusillus in error.

334 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1932. Hyperolius iisaramoae Loveridge, Proc. Biol. Soc. Washington, 45,

p. 63 (based on 5 ) : Mogogoni Swamp, south of Dar es Salaam,

Tanganyika Territory.
1935. Hyperolius inilnei Loveridge, Bull. Mus. Comp. Zool., 79. p. 18

(based on $ ) : Witu, Coast Province, Kenya Colony.
1935. Hyperolius translucens Power, Proc. Zool. Soc. London, p. 339,

figs. 6-7, pi. i, figs. 3-4 (based on $ and $ ) : Port St. Johns,

Pondoland, South Africa.
Range. Coastal *Keuya Colony ; *Tangaiiyika Territory ;
*Nyasaland ; *Natal and adjacent *Cape Province.

B. Species usually known only from the original descrip-
tion, fre(iuently based on a single individual. This list
is alphabetically arranged.

Hyperolius albifrons Ahl

1931. Hyperolius albifrons Ahl, Das Tierreich, no. 55, p. 354, fig. 230:

' ' Africa. ' '
Range. Mafia Island (//'rfe Parker :1937).

Hyperolius alfaofrenatus Ahl

1931. Hyperolius albofrenatus Ahl, Das Tierreich, no. 55, p. 315, fig.

189 (coll. Ule or Ulil) : "German East Africa."
Range. Tanganyika Territory.

Hyperolius albolabris Ahl

1931. Hyperolius albolabris Ahl, Das Tierreich, no. 55, p. 287, fig. 101
(coll. Neumann: 26.ii.1894) : Kwa Buosch or Bnorch, Tan
ganyika Territory.

Range. Tanganyika Territory.

Hyperolius bergeri Ahl

1931. Hyperolius bergeri Ahl, Das Tierreich, no. 55, p. 346, fig. 221 :

"Guaso" i.e. Uaso Narok, Keuj-a Colony.
Range. Kenya Colony.

Hyperolius glandicolor Peters

1878. Hyperolius glandieolor Peters, Monatsb. Akad. Wiss. Berlin, p.

208, pi. ii, fig. 9: "Taita," i.e. Teita, Kenya Colony.
Range. Kenya Colony.

Hyperolius guttolineatus Ahl

1931. Hyperolius guttolineatus Ahl, Das Tierreich, no. 55, p. 321, fig.

195 (coll. Uhl): "German East Africa."
19)1. HyperoUu.'^ vuinuorafus guttatoUneatus Laurent {lapsus for gtit-

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 335

tolineatus Ahl, though page and figure numbers are wrong and
should be as eited above), Ann. Sei. Roy. Zool. Belgique, 82.
p. 389.
Range. Tanganyika Territory.

Hyperolius mctmioratus campylogrammus Laurent '^^

19.'51. Hyperolius marmoratus campylogrammus Laurent, Ann. Soc. Roy.
Zool. Belgique, 82. p. 36: Bura, 1050 metres, Teita, Kenya
Colony.

Range. Kenya Colony.

Hyperolius quadrotomaculotus Ahl

1931. Hyperolius quadratomaculatus Ahl, Das Tierreich, no. 55, p. 412.

lig. il85 : Mohoro, Tanganyika Territory.
Range. Tanganyika Territor}^

Hyperolius scriptus Ahl

1931. Hyperolius scriptus Ahl, Das Tierreich, no. 55, p. 287, fig. 160:

Tanga, Tanganyika Territory.
Range. Tanganyika Territory.

Hyperolius tomieri Ahl

1931. Hyperolius tomieri Ahl, Das Tierreich, no. 55, p. 304, fig. 179:

I'kami, i.e. Uluguru Mountains, Tanganyika Territory.
Range. Tanganyika Territory.

Hyperolius viridiflavus pitmoni Laurent

1951. Hyperolius virieliflavus pitmani Laurent, Ann. Soc. Roy. Zool.

Belgique, 82, p. 394: Lake Bunyoni, southwest Uganda.
Range. Uganda.

Family RANIDAE^^""
Genu.s ARTHROLEPTIDES Nieden

1910. Arthroleptides Nieden, Sitzb. Ges. Naturf. Freunde Berlin, p. 445.
Type by monotypy : A. martiensseni Nieden.

Arthroleptides martiensseni Nieden
I^sambara Montane-torrent-Frog

245 To which Laurent wonhl assign the Mount Mbololo frog that I refer with
misg^ivings to flavoyuttatua .\hl.

246 This is eniploj-pd in its generall.v accepted sense. Laurent (1951, Revue
Zool. Bot. Afr.. 45, p. 120) has proposed radical changes in which he would
transfer Arthrolepth and Hrmi!<us to a new concept, the HYPEROLIIDAE.

336 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

1910. Arthroleptides martiensseni Nieden, Sitzb. Ges. Naturf. Freunde
Berlin, p. 445: Amani, Usambara Mountains, Tanganyika Terri-
tory.
Range. Rocky montane streams of Usambara, Magrotto and
Uluguru Montains, Tanganyika Territory.

Arthroleptides dutoiti Loveridge Elgon Montane-torrent-Frog
1935. Arthroleptides dutoiti Loveridge, Bull. Mus. Comp. Zool., 79. p. 17:

Koitobos Eiver, 7200 feet, Mount Elgon, Kenya Colony.
Range. Rocky montane streams of Mount Elgon, Kenya Col-
ony.

Genus RANA Linnaeus

1758. FMiia Linnaeus, Syst. Nat., ed. 10, 1. p. 210. Type by designation
of Fitzinger in 1843 : B. temporaria Linnaeus.

1814. Eanaria Eafinesque, Specchio Sci. (Palermo), 2, No. 10, p. 102.
Substitute name for Eana Linnaeus.

1838. Hylarana^*' Tschudi, Class. Batr. Eept. (reprinted 1839), p. 78.
Type by monotypy : Eyla erytJiraea Schlegel.

1838. Strongylopiis Tschudi, Class. Batr. Eept. (reprinted 1839), p. 79.
Type by monotypy: S. fasciatus Tschudi.

1838. Fyxicephalus Tschudi, Class. Batr. Eept. (reprinted 1839), p. 83.
Type by designation of Boulenger in 1920: P. adspers^is
Tschudi.

1841. Limnodytes Dumeril & Bibron, Erp6t. Gen., 8, p. 510. Substitute
name for Hylarana Tschudi.

1843. Pelophylax Fitzinger, Syst. Eept., p. 31. Type by original designa-
tion : Eana escidenta Linnaeus.

1843. Limnophiius Fitzinger (not of Agassiz:1842), Syst. Eept., p. 31.
Type by original designation : Eana mascareniensis Dumeril &
Bibron.

1843. Euphlyctis Fitzinger, Syst. Eept., p. 31. Type by original designa-
tion: Eana Icsdlienaultii Dumeril & Bibron = Eana cyano-
phlyotis Schneider.

1843. Phrynoderma Fitzinger, Syst. Eept., p. 31. Type by original
designation: EaTia cuiipora Dumeril & Bibron = Eana hexa
dactyla Lesson.

1843. Eydrophylax Fitzinger, Syst. Eept., p. 31. Type by original desig-
nation: Eana malabarica Tschudi.

1843. Limnonectes Fitzinger, Syst. Eept., p. 31. Type by original desig-
nation: Eana Tcuhlii Dumeril & Bibron.

247 Amended to Hjflorana by various authors.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 337

1858. Tomopterna Giinther, Cat. Batr. Sal. Brit. Mus., p. 7. Type by

designation of Boulenger in 1920: Pyxioephalus delalandii
Tschudi.
18.58. Sphaerotheca Giinther, Cat. Batr. Sal. Brit. Mus., p. 20. Type by
monotypy: S. sirigata Giinther = Rana. hreviceps Schneider.

1859. Ranula Peters, Monatsb. Akad. Wiss. Berlin, p. 402. Type by

monotypy: R. gollmerii Peters == Roma palmipes Spix.

1860. Dicroglossus Giinther, Proc. Zool. Soc. London, p. 158. Type by

monotypy: D. adolfi Giinther.
1863. Hoplohatrachus Peters, Monatsb. Akad. Wiss. Berlin, p. 449. Type

by monotypy: H. ceylanicvs Peters.
1865. Amolops Cope, Nat. Hist. Eeview, 5. p. 117. Type by monotypy:

Polypcdates afghanus Giinther.
18(57. Pohlia Steindachner, Amphibia, in Eeise osterr. Fregatte Novara,

p. 15. Type by monotypy: Rama palmipes Spix.
1868. Trypheropsis Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 117.

Type by original designation: T. chrysoprasinus Cope.

1868. Pachrjbat radius iMivart (not of Kefferstein:1868), Proc. Zool.

Soc. London, p. 559. Type by monotypy: P. robustus Mivart.

1869. Clinotarsus Mivart, Proc. Zool. Soc. London, p. 227. New name

for Pachybatrachus Mivart (preoccupied).
1881. Maltzania Boettger, Abhand. Seuckeuberg. Naturf. Ges., 12, p.

417. Type by monotypy: il. bufonia Boettger.
1891. Crotaphitifs Schulze, Jahresb. Naturw. Yer. Magdeburg, 189U,

p. 176. Type by designation of Stejneger in 1907: Rarui iem-

poraria Linnaeus.
1891. Baliopygus Schulze, Jahresb. Naturw. Ver. Magdeburg, for 1890,

p. 177. Type by designation of Stejneger in 1907: Rana escu-

lenta Linnaeus.

1893. Phrynopsis Pfeffer, Jahrb. Hamburg. Wiss. Anst., 1892, 10. p. lUl.

Type by monotypy: P. boidcngerii Pfeffer ^ juv. Pyxicephalus
edulis Peters.

1894. Levirana Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 197. Type

by original designation: L. vibicaria Cope.
1907. HildebrandUa Nieden, Sitzb. Ges. Naturf. Freunde Berlin, p. 229.

Type by monotypy: Pyxicephalus oniatus Peters.
1917. Aubria Boulenger, C. E. Acad. Sci. (Paris), 165. p. 988, footnote.

Type by monotypy: Rana subsigillata A. Dumeril.
1920. Ptychadena Boulenger, Eec. Indian Mus., 20. p. 5. Type by original

designation: Rana mascaretiiensis Dumeril & Bibron.
1924. Ranosoma Ahl, Aroh. Naturg.. 90. Abt. A, p. 250. Type by

monotypy: R. schereri Ahl = Rana occipitalis Giinther.
1927. Pseudoxenopus Barbour & Loveridge, Proc. New England Zool.

338 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Club, 10, p. 14. Type by original designation: P. alleni Barbour
& Loveridge.-^^

1931. Ahraria Parker, Proc. Zool. Soc. London, 1930, p. 898. Type by
monotypy: A. cotti Parker = Bana floweri Boulenger.

1931. Palcorana Scortecci, Atti. Soc. Ital. Sci. Nat. (Milano), 70. p. 17.
Type by designation of Parker in 1936: Eana beccarii Boulen-
ger.

1931. Gigantorana-*^ Noble, Biology of the Amphibia (New York), p.
159. Type by monotypy: Bana goliath Boulenger.

Rona albolabris albolabris Hallowell
Disk-toed White-lipped Frog
1856. Baiia albolabris Hallowell, Proc. Acad. Nat. Sci. Philadelphia, p.

153.
1903. Chlromantis lepus Andersson, Verh. Zool. Bot. Ges. Wien, 53, p.

142 (53.5 mm. ex.) : Cameroon.^"
Range. Southern *Sudan; Uganda; western Tanganyika Ter-
ritory (Bukoba) ; west to Angola (where it meets with the race
a. parkeriana Mertens), north to Cameroon, west to Portuguese

ri * 251

Guinea.

Rccna galamensis bravana (Peters)

1882. Limnodytcs hmvanus Peters, Sitzb. Ges. Naturf. Freunde Berlin,

p. 3: "Barawa, " i.e. Brava, Somalia.
1929. Buna magrelti Srortecei, Atti. Soc. Ital. Sci. Nat. (Milano), 68.

p. 182, pi. ix, figs. 1-3: Ghinda, Eritrea.
1929. Bana fiechteri Scortecci, Atti. Soc. Ital. Sci. Nat. (Milano), 68.

p. 248, pi. xii, figs. 2-4: Villaggio Duca degli Abruzzi, Somalia.
1931. Bana somalica Scortecci, Atti Soc. Ital. Sci. Nat. (Milano), 69,

p. 320: Villaggio Duca degli Abruzzi, Somalia.
Range. Somalia; coastal *Kenya Colony; ^Tanganyika Terri-
tory ; *Pemba Island ; *Zanzibar Island ; Mozambique ; Nyasa-
land; Northern Rhodesia; eastern *Belgian Congo.

Rana galamensis galamensis Dumeril & Bibroii

248 Based on a subadult without vomerine teeth; the frog is now known as
Rana crassipes alleni Barbour & Loveridge.

2i9 Apparently attributed to Nieden in error.

230 c. leptus is unquestionably a synonym as may be seen from tbe figure sub-
sequently (1905) published bv Andersson. Less certain was Noble's action in
synonym'izing R. zenkeri Nieden, 1908, Mitt. Zool. Mus. Berlin, 3. p. 497 (92
mm. adult and 8 others), from Bipiudi. Cameroon, for it appears recognizable.
The Museum of Comparative Zoology has both from Metet, Cameroon.

231 Of this common arlioreal frog tlie Museum has examples from Siulan ;
Belgian and French Congo ; French Cameroon : Nigeria : Ivory Coast ; Liberia ;
Sierra I.,eone.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 339

1841. Bana galamensis Dumeril & Bibron, Eipet. Gen., 8, p. 367: Galam

Lakes, Senegal.
ISOfi. Hana oiihanrjhiensis Mocquard, C. R. Soc. Pliilom. Paris, No. 19,
p. 234: "Upper Oubanghi, " i.e. Oubangui Eiver, French Equa
torial Africa.
Range. *Sudan ; *Uganda ; Belgian Congo ; Belgian Ruanda-
Urundi ; French and British Cameroon ; Nigeria ; *Gold Coast :
French Guinea; Portuguese Guinea; *Gambia; Senegal.

Rana wittei (Angel)

1924. Phrynoiatracliu^ Wittei Angel, Bull. Mus. Hist. Nat. (Paris), 30,

p. 130: Molo, Mau Escarpment, Kenya Colony.

1925. Bana aherdariensis Angel, Eept. et Batr., in Voyage de Ch.

Alluaud et R. Jeannel en Afrique Orientale (1911-1912).
Resultats scientifiqiies. Yertebrata. (Paris), 2. p. 42, pi. ii, figs.
1-2: Mount Kinangop, Aberdare Mountains, Kenya Colony.
Range. Montane marshes of Ethiopia {iide Angel) ; *Kenya
Colony and eastern *Belgian Congo.

Rana fuscigula chapini Noble

Montane-forest-stream Dusky-throated-Frog
1924. Bana chapini Noble, Bull. Amer. Mus. Nat. Hist., 49. p. 214, fig.

6a: Batatna, Stanleyville district, Belgian Congo.
Range. Forest streams of ^Ethiopia; *Uganda; *Kenya Col-
ony; ^Tanganyika Territory and *Belgian Congo.

Rana fuscigula angolensis Boeage"'''

Savanna-stream Dusky-throated-Frog
1866. Bana angoJensis Bocage, .Torn. Sci. Lisboa, 1. p. 73: Duque de
Braganca, Angola.

1893. Bana myassae Giinther, Proc. Zool. Soc. London, 1892, pp. 555, 558:

Shire Highlands, Nyasaland.

1894. Bana queeketti Boulenger, Proc. Zool. Soc. London, p. 643,^^^ pi.

xxxix, fig. 1: Pietermaritzburg, Natal.
1896. Bann Xutti Boulenger, Ann. Mag. Nat. Hist., (6) 18. p. 467:

Lake Tanganyika.
Range. *Uganda ; *Kenya Colony ; *Tanganyika Territory :

252 Possibly not separable from /. fiiscir/ula from which it is distinguished only
by its longer hind legs, a character which it shares with /. chapini.

•-■">3 Though qtivckrtti was svuonymi/.ed liy Boulenger himself with /. fuscigula,
with which it appears to agree in limb length. Dr. V. F. PitzSimons informs me
ili2.x.5«» that it does not in the least resemble typical fuscigula and should, in
bis opinion, be assigned to the synonymy of /. angolensis. He regards /. fuscigula
as being mure or less confined to the Cape ; unfortunately short-linibed individu-
als are present throughout much of the range of /. nngolensis.

340 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Mozambique ; *Nyasaland ; Northern and Southern Rhodesia ;
Bechuanaland ; *Transvaal; *Natal; *Orange River; *Cape
Province ; *Angola ; *Belgian Congo ; *Belgian Ruanda-Urundi.

Rana fuscigula fuscigula Dumeril & Bibrou
Short-limbed Dusky-throated-Frog

1841. Eajia fuscigula Dumeril & Bibron, Erpet. Gen., 8. p. 386: Cape of
Good Hope, i.e. South Africa.

Range. Owing to long confusion with E. f. chapini and B. f.
angoJensis little reliance can be placed on records in the litera-
ture. I have examined the material on which those from Kenya
Colony and Tanganyika Territory were based and found them
to be R. f. chapini. I have collected frogs conforming to /. fusci-
gula at MiLshongero, Lake Mutanda, southwest *Uganda (which
may be an area of intermediates as /. angolensis was also pres-
ent there); *Nyasaland; *Bechuanaland Protectorate; *Orange
River; *Cape Province.

Rana christyi Boulenger

1919. Bana christyi Boulenger, Revue Zool. Afr., 7. p. 5: "Madie, "

i.e. Medje, Ituri, Belgian Congo.
Range. *Uganda and *Belgian Congo.

Rana oxyrhynchus gribingviiensis Angel
Montane-forest Sharp-nosed-Prog
1922. Bana (Ptychadena) Grihinguiensis Angel, Bull. Mus. Hist. Nat.
(Paris), 28. p. 399, fig. — : Fort Crampel, Lake Chad, French
Equatorial Africa.
Range. Streams in montane forest, or recently deforested
areas, of *Uganda; *Kenya Colony; *Tanganyika Territory;
*Nyasaland ; *Belgian Congo ; French Congo ; French Equatorial
Africa ; *French Cameroon ; Nigeria ; Ivory Coast ; *Liberia.

Rana oxyrhynchus oxyrhynchus A. Smith
Savanna Sharp-nosed-Frog
1849. Bana oxyrhynchus "Sundevall" A. Smith, Illus. Zool. S. Africa,
Kept., pi. Ixxvii, figs. 2 and 2a-2c : Kaffirland and the region of
Port Natal, i.e. Durban, Natal.
1868. Bana anchietae Bocage, Proc. Zool. See. London, 1867, p. 843,

fig. 1 (48 mm. $ ) : Benguela, Angola.
1897. Phrynobatrachns hailensis Meek, Field Mus. Nat. Hist. Zool. Series,
1, p. 175: "Haili," i.e. Halleh, British Somaliland.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 341

1906. Sana Theileri Mocquard, Bull. Mus. Hist. Nat. (Paris), 12. p. 252:
Nelspruit, Transvaal.

1923. Ptychadena aberae Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p.
97: Abera, near "Jandjan" (I Jiran), southwest Ethiopia.

1933. Bana oxyrliynchus migiurtina Scortecei, Atti Soc. Ital. Sci. Nat.
(Milano), 72. pp. 12, 51, pi. i, fig. 3: Uadi Arro, Somalia.

Range. Chiefly in stream^s of lowland or plateau savanna in
the *Sudan, Eritrea, *Ethiopia, French and *British Somaliland,
Somalia, *Uganda Protectorate, *Kenya Colony, ^Tanganyika
Territory, *Pemba, Zanzibar and Mafia Islands, *Mozambique,
*Nyasaland, Northern and *Southern Rhodesia, Bechuanaland
Protectorate, *Transvaal, *Natal, *Cape Province, Southwest
Africa, Cabinda, Angola, Belgian Congo and *Belgian Ruanda-
Urundi. Reported from almost every country in Africa south of
15° N., but many western records may have resulted from con-
fusion with the montane forest race listed above or with R. o.
superciliaris Giinther of Sierra Leone.

Rana floweri Boulenger-^*

1917. Bana -floweri Boulenger, Ann. Mag. Nat. Hist., (8) 20. p. 417:

Rosaires, Blue Nile, Sudan.
1923. Bana erlangeri Ahl, Mitt. Zool. Mus. Berlin, 11, p. 5: Lake Abaya,

northeast of Lake Stephanie, Ethiopia.
1925. Bana harhouri Loveridge, Proc. Zool. Soc. London, p. 776, pi. i.

fig. 2; Nyambita, Mwanza, Tanganyika Territory.
1931. Ahrana cotti Parker, Proc. Zool. Soc. London, 1930, p. 898, fig. 1:

Charre, Mozambique.
Range. *Egypt (Giza: M.C.Z. 3138-40), Sudan, *Kenya Col-
ony, *Tanganyika Territory, *Mozambique. (I regard the French
Cameroon record of Monard, 1951, confirmed by Laurent, as
somewhat questionable.)

Rana mascoreniensis mascareniensis "^^ Dumeril & Bibron
Common Mascarene-Frog

254 For long confused with o. oxyrhynchus, from which floweri may be dis-
tinguished by the fine black vermiculatlons on its buttocks and shorter hindllmb,
whose tibio-tarsal articulation, when directed forwards, reaches only to the eye
or nostril (in o. oxyrhynchus to end of snout, usually far beyond).

255 Unless a $ (M.C.Z. 28622) from Liwale, southeastern Tanganyika Terri-
tory is correctly assigned, so far as I am aware the perfectly valid species or
subspecies R. mascareniensis mossambica Peters (1854), together with its
synonyms trinotlis var. Bottgeri Pfefifer (1893) and vernayi FitzSimons (1932)
does not occur in the area covered by this list. However, early records from
British East Africa of the Senegalese species trinodis Boettger (1881), of which
schubotzi Sternfeld (1917) is a synonym, may have been based on mossambica'!

342 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

1841. Eana Mascarcniensis Dumeril & Bibron, Erpet. Gen., 8. p. 350:

Madagascar; Mauritius; Seychelles.
1855. Eana nilotica Seetzen, Eeise durch Syrian, Palast., AegjT)t, etc.,

3, p. 490 : Egypt.
1857. Eana savignyi Jan, Cenni. Mus. Civ. Milano, p. 52 (probably

Egypt, but this work is not available to me).
1864. ■^^Eama idae Steindachner, Verb. Zool.-Bot. Ges. Wien, 14. p. 266,

pi. xii, fig. 1 : Madagascar.

1864. lEana nigrescens Steindachner, Verb. Zool.-Bot. Ges. Wien, 14. p.

268, pi. xii, figs. 2 and 2a-2c : Madagascar.

1865. Eana spinidactyla Cope, Proc. Acad. Nat. Sci. Philadelphia, p.

197: Natal.
1881. Eana ahyssinica Peters, Sitzb. Ges. Naturf. Freunde Berlin, p.

163: Ailet near Massawa; and Keren, Bogos, Eritrea.
1886. Eana Neivtanii Bocage, Jorn. Sci. Lisboa, 11, pp. 70, 73: Sao

Tome, Gulf of Guinea.
Range. *Egypt ; *Sudan; *Etliiopia; * Uganda; *Kenya Col-
on}^ ; *Tanganyika Territory ; *Zanzil)ar and Mafia Islands ;
*Madagascar : *Mozambique ; *Nyasaland ; *Northern and
^Southern Rhodesia; *Transvaal; *Angola ; *French Congo;
*Nigeria; *Gold Coast; *Ivory Coast; *Liberia ; *Sierra Leone.

Besides which it has been reported from almost every country
in Africa, but due to confusion with closely related forms I have
listed only those countries represented in the collection of the
Museum of Comparative Zoology.

Rana mascareniensis venusta Werner
Primary-forest Mascarene-Frog
1908. Eana vcnusia Werner, Sitzb. Akad. Wiss. Wien, 1907, 116, Abt. 1.
pp. 1889, 1892, pi. iv, fig. 11: Mongala, Sudan; Entebbe.
Uganda ; Lagos, Nigeria.
Range. Primary or gallery forested areas of the southern
Sudan; *Uganda; *Kenya Colonj^; *Tanganyika Territory, west
through *Belgian Ruanda-Urundi and *P)elgian Congo; *French
and British Cameroon to Lagos, Nigeria.

Rana mascareniensis uzungwensis LoAcridge
Montane Mascarene-Frog

1932. Eana mascareniensis u:;ungivensis Loveridge, Bull. Mus. Comp.

Zool., 72, p. 384: Dabaga, Uzungwe Mountains, Tanganyika

Territory.
1952. Ptycliadena macrncepliala Laurent, Revue Zool. Bot. Afr., 46, ii.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 343

22: Upper Lubitshako River, 1900-2000 metres, south Kivu, Fizi

Territory, Belgian Congo.
Range. Upland marshes of southern *Tanganyika Territory ;
*N3'asaland ; *Southeru Rhodesia ; *Angola ; Belgian Congo and
Belgian Ruanda-Urundi.

Rana loveridgei ( Laurent )"^^

1954. Piychadena loveridgei Laurent, Ann. Mus. Roy. Congo Beige, 34,
p. 14, pi. i, fig. 4; pi. ii, fig. 1; pi. iii, figs. 3-4: Tare, Busanza,
ca. 1800 metres, Astrida region, Belgian Ruanda-Urundi.
Range. South-west Uganda; *Tangauyika Territory; Belgian
Ruanda-Urundi ; Belgian Congo ; Angola.

Rana ansor(^i Boulenger

1905. Eana ansorgii Boulenger, Ann. Mag. Nat. Hist., (7) 16. p. 107,
pi. iv, fig. 1 : Between Benguela and Bihe, Angola.

Range. Southwest *Tanganyika Territory ; Mozambique ; *Ny-
asaland; *Northern Rhodesia; * Angola north to Lower Congo
River and Katanga, Belgian Congo.

Rana fasciata merumontana Lonnberg

Northern Tanganyika Striped Grass-Frog
1907. Rana merinnontana Lonnberg, in Sjostedt, Wiss. Ergeb. Schwed.
Zool. Exped. Kilimandjaro, Meru umgeb, Massaisteppen, No. 4,
p. 21, pi. i, figs. 4a-b: Mount Meru, Tanganyika Territory.
Range. Montane meadows of northern *Tanganyika Terri-
tory.

Rana fasciata fiillebomi Nieden

Southern Tanganyika Striped Grass-Frog
1910. Rana fiillehorni Nieden, Sitzb. Ges. Naturf. Freunde Berlin, p.
436: Crater Lake, Ngosi Volcano, Poroto Mountains, Tan-
ganyika Territory.

Range. Montane meadows of southern *Tanganyika Terri-
tory and *Nyasaland.

Rana stenocephala Boulenger Uganda Striped Grass-Frog
1901. Rana stenocepliala Boulenger, Ann. Mag. Nat. Hist., (7) 8, p. 515:

Entebbe, Uganda.
Range. *L'ganda (known only from the type locality).

256 Whether this is a valid "species" remains to be seen ; it is based in part on
small ^ paratypes of R. m. uzungwcnsiH Loveridge (see above) from Dabaga and

Kigogo, T. T. However, Laurent had abundant material and the countries given
in the range are taken from his paratypes.

344 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Rcrna occipitalis Giinther Groove-crowned Bullfrog

1858. Bana occipitalis Giinther, Cat. Batr. Sal. Brit. Mus., p. 130, pi. xi:
Gambia ; West Africa ; Africa.

1863. Ban^ hydraletis "Boie" Peters (nomeJi nudum), Monatsb. Akad.

Wiss. Berlin, p. 78 : Type in Leiden Museum examined by
Peters.

1864. Bona bragantina Bocage, Revue Mag. Zool. (2) 16, p. 253:

Duque de Braganca, Angola.

1924. Banosoma sehereri AM, Arch. Naturg., 90, Abt. A, p. 250 : Senegal.

1925. Bana miuanzae Loveridge, Proc. Zool. Soc. London, p. 772, pi. ii:

Simiyu River at Sagayo, Mwanza Province, Tanganyika Terri-
tory.
1954. Bana accidentalis Lamotte & Zuber-Vogeli (lapsus for occipitalis

Giinther), Bull. Inst. Franc. Afr. Noire, 16, Ser. A, p. 940.
Range. Reported from French Morocco ; Algeria ; Libya ;
*Sudan; Eritrea; *Uganda; Kenya Colony; *Tanganyika Terri-
tory ; ^Northern Rhodesia ; Angola ; Cabinda ; *Belgian and
French Congo ; French Equatorial Africa ; Sao Thome ; Fer-
nando Po ; *French and British Cameroon ; Dahomey ; Togo ;
*(jlold Coast; *Liberia; *Sierra Leone; French and Portuguese
Guinea ; *Gambia ; *Senegal.

Rana adspersa edvdis (Peters) Eastern Burrowing-Bullfrog
1854. Pyxicephnlus edulis Peters, Monatsb. Akad. Wiss. Berlin, p. 626:

Boror ; Tete ; and Mozambique Island, Mozambique.
1893. Phrynopsis Boiilengerii Pfefifer, Jahrb. Hamburg. Wiss. Anst.,

1892, 10. p. 101, pi. ii, figs. 5-7: Mozambique.
1916. Pyxicephalus flavigida Calabresi, Monit. Zool. Ital. (Firenze), 27,

p. 34, pi. ii, fig. 1 : Oroflillo, Somalia.
1924. Phrynopsis usambarae Ahl, Zool. Anz., 60, p. 271: Usambara,

Tanganyika Territory.
1927. ? Pyxicephalus obbianus Calabresi, Atti Soc. Ital. Sci. Nat. (Milano),

66, pp. 15, 35, pi. i, fig. 1: Dolobseio, near Obbia, Somalia.
Range. Somalia, south through *Kenya Colony ; *Tanganyika
Territory ; *Mozambique ; Nyasaland and Northern Rhodesia to
the Zambezi River. ^'''

Rana delalandii delalandii (Dumeril & Bibron)

Delaland 's Burrowing-Frog

1841. Pyxicephalus Delalandii Dumeril & Bibron, Erpet. Gen., 8. p. 445,

pi. Ixxxvii, figs. 1 and lab : South Africa.

257 For detailed locality records, see Loveridge, 1950, Journ. E. Afr. Nat. Hist.
Soc. for 1947-1948, 19, pp. 253-255. The Museum of Comparative Zoology has
typical It. a. adspersa (Tschudi) from Southern Rhodesia and the Transvaal,
south to Cape Province.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 345

1854. Pyxicephalus marmoratus Peters, Monatsb. Akad. Wiss. Berlin,

p. 626: Boror, Mozambique.
1884. Phrynobatrachus monticola Fischer, Jahrb. Hamburg. Wiss. Anst.,

1, p. 26: Southern Uaso Nyiro, Masailand, Kenya Colony.
1927. Eana (Tomopterna) hieroglyphica Ahl, Sitzb. Ges. Naturf.

Freuude Berlin, p. 42: So. Omadu, Somalia.
1936. Arthroleptis agadezi Angel, Bull, Soc. Zool. France, 61. p. 275:

Agadez, Air, Sahara, French West Africa.
1938. Arthroleptis ahli Deckert, Sitzb. Ges. Naturf. Freunde Berlin,
pp. 127, 168, fig. 46: Lichtenstein (approx. 32°50' S., 26°E.,
between Bedford and Cheviot Falls, eastern Cape Province),
South Africa.
1944. Arthroleptis rosei Hoffman, Soolog. Navors. Nas. Mus., Bloem-
fontein, 1. p. 174, figs. 1-4: "Chitiala," i.e. Chitala River,
Nyasaland.
Range. Arid savanna regions in French West Africa (at
Agadez: fide Guibe), west through the Sudan to Eritrea; south
through *Ethiopia; *British Somaliland; *Somalia; Uganda;
*Keuya Colony ; *Tanganyika Territory ; Mozambique ; *South-
ern Rhodesia ; Bechuanaland ; *Transvaal ; *Orange River ;
*Cape Province to Southwest Africa where it meets with the
race R. (]. cryptotis Boulenger.

Rona tuberculosa (Giinther)-^** Angolan Burro wing-Frog

1864. Pyxicephalus tuberculosa Giinther, Proc. Zool. Soc. London, p.

479, pi. xxxiii, fig. 1 : Pungo Ndongo, Angola.
1896. Rana pulchra Boulenger, Ann. Mag. Nat. Hist., (6) 18. p. 468:

Lake Tanganyika (without indication as to which coast).
Range. ''Lake Tanganyika;" Angola.

Rana ornata ornata (Peters)

Northern Ornate Burrowing-Frog
1878. Pyxicephalus ornatus Peters, Monatsb. Akad. Wiss. Berlin, p. 207,

pi. ii, fig. 7: "Taita," i.e. Teita, Kenya Colony.
1912. Eana macrotym'paniim Boulenger, Ann. Mag. Nat. Hist., (8) 10,

p. 140: West of Juba River, Gallaland, Ethiopia.
1919. Eana (Hildebrandtia) togoensis Boulenger,-^" Trans. Roy. Soc.

S. Africa, 8, p. 34: Mangu, Togo.
Range. Somalia; Kenya Colony; *Tanganyika Territory;

-5S Miss A. G. C. Grandison informs me (29.xi.56) that Boulenger himself
synonymized pulchra with tuberculosa — of which we have only Angolan ma-
terial. In details our frogs do not agree too well with the description of pulchra,
and the relationship with delalandii requires investigation.

259 Kew name for H. omatisslma Nieden (not of Bocag^), proposed by Boulen-
ger, sight unseen.

346 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

*BeIgian Congo; French Equatorial Africa (det. A.L.) ; Togo;
*CT0ld Coast.

Genus PHRYNOBATRACHUS Glinther

260

1849. Stenorhynchus A. Smith (not of Hemprich: 1820), Illus. Zool. S.
Africa, Eept., App., p. 23. Type by monotypy: S. natalensis
Smith.

1862. Phrynobatrachus Giinther, Proc. Zool. Soc. London, p. 190. Type

by monotypy: P. natalensis Giinther (correct).

1863. Leptoparius Peters, Monatsb. Akad. Wiss. Berlin, p. 452. New

name for StenorhyncJnis A. Smith (preoccupied).

1925. Hylarthroleptis Ahl, Sitzb. Ges. Naturf. Freunde Berlin, 1923,

p. 98. Type by present designation: H. accraensis Ahl.

1926. MicrobatracJius Hewitt (not of Koux:1910), Ann. S. African Mus.,

20, p. 420. Type by original designation: Phrynobatrachus
capensis Boulenger.
1926. Microbatrachella Hewitt, Ann. S. African Mus., 20, p. 420, cor-
rigenda slip. Xew name for Microbatrachus Hewitt (preoccu-
pied).

Phrynobatrachus perpahnatus Boulenger

1898. Phrynobatrachus perpalmatu.'i Boulenger, Proc. Zool. Soc. London,

p. 479, pi. xxxviii, fig. 1 : Lake Mweru, Northern Ehodesia.
1924. Phrynobatrachus perpahnatus werneri Ahl, Zool. Anz., 60. p. 273:

El Grassi, etc., Sudan.
Range. Sudan ; *Tanganyika Territory ; ^Mozambique ; *Ny-
asaland ; *Northern Rhodesia ; *Belgian Congo.

Phrynobatrachus acridoides (Cope)

1867. Staurois acridoides Cope, Journ. Acad. Nat. Sci. Philadelphia,

6, p. 198 : Zanzibar Island.
1919. PhrynoboArachus Boulengeri Witte, Revue Zool. Afr., 6, p. 225:

Beira and Coguna, Mozambique.
1923. Hylarthroleptis medilineatus Ahl, Sitzb. Ges. Naturf. Freunde

Berlin, p. 100: " Tscharra, Dauafluss," i.e. Charra, Tana River.

Kenya Colony.
1923. Hylarthroleptis janenschi Ahl, Sitzb. Ges. Naturf. Freunde Berlin,

p. 100: Tendaguru, near Lindi, Tanganyika Territory.

Range. Sudan ; Eritrea ; Somalia ; *Kenya Colony ; *Tan-

260 Despite Hewitt's defense (1926, Ann. S. African Mus., 20, p. 425) of the
generic status of Natalohatrachua (Hewitt & Methueu, 1913, Trans. Roy. Soc. S.
Africa, 3, p. 107. Type by monotypy N. boncberyi Hewitt & Metliuen) it should
probably be regarded as a subgenus of Phrynohutrackun of which Noble regarded
it as a synonym.

LOVERIDQE: E. AFRICAN REPTILES AND AMPHIBIANS 347

ganyika Territorj' ;~^^ Pemba, Zanzibar and Mafia Islands;
Mozambique ; Northern Rhodesia ; French Equatorial Africa
(fide Sternfeld:1917) ; *French and British Cameroon; also re-
ported from Gold Coast (Witte :1919) ; French Guinea (Klap-
tocz:191;3; Chabanaud:1921) ; Gambia (Witte :1919) ; Senegal
(Mertens:1938).

Phrynobcrtrachus pakenhami Loveridge

1941. Phnjnohatrachus i^akenlMmi Loveridge, Proc. Biol. Soc. Washing-
ton, 54. p. 178: Machengwe Swamp, near Wete, Pemba Island.
Range. Pemba Island.

Phrynobcrtrachus kinangopensis Angel

1924. Fhrynobairachns Kinangopensis Angel, Bull. Mus. Hist. Nat.
(Paris), 30, p. 130: Mount Kinangop, Aberdare Mountains,
Kenya Colony.
Range. From 5000 (at Nairobi) to 11,000 feet (on Mount
Kenya), in highlands of Kenya Colony.

Phrynobatrachus plicatus (Giinther)

1858. Byperoliu.s plicatus Giinther, Cat. Batr. Sal. Brit. Mus., p. 88, pi.

vii, fig. C: Coast of Guinea.
1900. Phryjiobat radius auritus Boulenger, Proc. Zool. Soc. London, 2,

p. 440, pi. xxviii, fig. 2: Benito Eiver, French Congo.
1919. Phrynobatrachus discodactylus Boulenger, Revue Zool. Afr., 7,

p. 7: "Madie," i.e. Medje, Ituri, Belgian Congo.
Range. *Ilganda (Budongo Forest) and Belgian Ruanda-
Urundi, west through *Belgian Congo to Cabinda, north and
west through French Congo ; Rio Muni ; Fernando Po ; *French
and British Cameroon; Togo; Gold Coast; *Liberia to *Sierra
Leone.

Phrynobatrachus krefftii Boulenger

1909. Phrynobatrachus krefftii Boulenger, Ann. Mag. Nat. Hist., (8) 4,
p. 49t> : Amani, Usambara Mountains, Tanganyika Territory.

Range. Usambara and Magrotto Mountains, Tanganyika Ter-
ritory.-®"

261 Frogs from the Uluguru Mountains of Tanganyika, referred with a query
to ogoensis Boulenger by P.arbour and Loveridge (1928), are referable to acri-
doidea. The two species are so similar that one is inclined to doubt all West
.\frican records of "acridoidcs."

2G2 Progs from Kuwenzori and Kuanda-Urundi referred to krefftii by Nieden
(1915) were subsequently renamed versicolor by Ahl (1924).

348 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Phxynobotrachus dendrobates (Boulenger)

1919. Arthrolepiis dendrohates Boulenger, Revue Zool. Afr., 7, p. 8:

"Madie, " i.e. Medje, Ituri Forest, Belgian Congo.
1924. Phrynohatrachus pctropedetoides Ahl, Zool. Anz., 61, p. 102:
Ruwenzori Mountains and west of Lake Edward, Belgian Congo.
Range. *Uganda ; Belgian Riianda-Urundi and the *Belgian
Congo.-"'

Phrynobcrtrachus versicolor Ahl

1924. Phrynobatrachiis versicolor Ahl, Zool. Anz., 61. p. 100: Ivugege
Forest, Belgian Ruanda-Urundi.

Range. *Uganda ; *Belgian Ruanda-Urundi and the *Belgian
Congo.

Phrynobatrachus graueri (Nieden)

1910. Arthrolcptis graueri Nieden, Sitzb. Ges. Naturf. Freunde Berlin,
p. 441 : Rugege Forest, Belgian Ruanda-Urundi.

Range. Western *Kenya Colony ^*^* and *Uganda, west through
*Belgian Ruanda-Urundi to the *Belgian Congo.

Phrynobatrachiis natalensis (Smith)

1849. Stcrnorhynclius natalensis A. Smith, Illus. Zool. S. Africa, Rept.,

App., p. 24: Natal, South Africa.
1862. Phrynobatrachus ■natalensis Giinther, Proc. Zool. Soc. London, p.

190 : Natal, South Africa.
]8()2. Bicroglossus angustirostris Cope, Proc. Acad. Nat. Sci. Philadel-
phia, p. 341: Umvoti, Natal (by inference only).
1875. Arthrolcptis natalensis var. irrorata Peters, Monatsb. Akad. Wiss.

Berlin, p. 210: Accra, Gold Coast.
1894. Phrynobatrachus ranoides Boulenger, Proc. Zool. Soc. London, pp.

641, 644, pi. xxxix, fig. 2 : Pietermaritzburg, Natal.

19U4. Phrynobatrachus natalensis fonna gracilis Andersson, Results

Swed. Exped. Zool. to Egypt and the White Nile 1901, \. No. 4,

p. 10, figs. — : Ghrab el Aish, south of Kaka on the White

Nile, Sudan.

Range. *Sudan; Ethiopia; *Uganda; *Kenya Colony; *Tan-

ganyika Territory ; Zanzibar Island ; Mozambique ; *Nyasaland ;

*Northern and *Southern Rhodesia; Bechuanaland ; *Transvaal ;

*Xatal; Orange River; *Cape Province; Southwest Africa;

*Angola; *Belgian Congo; Belgian Ruanda-Urundi; French

263 The record from Accra, Gold Coast, of Deckert (1938), certainly requires
verilicatiou.

-64 That is, Kaimosi and the Yala River region of Kakamega ; the Mount
Kinangop record of Angel (1925) is unacceptable.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 349

Cameroon; Dahomey; Togo; *Liberia; *Sierra Leone; French
Guinea; Portuguese Guinea; *Gambia; French West Africa.

Phrynobcrtrachns keniensis Barbour & Loveridge

1928. Phrynobatrachus Tceniensis Barbour & Loveridge, Proc. New Eng-
land Zool. Club, 10, p. 89: Mount Kenya, Kenya Colony.
Range. Upland meadows (Kikuyu; Molo; Mt. Kinangop ; Mt.

Kenya and Uplands) of Kenya Colony.

[Phrynobatrachus bottegi (Boulenger)]

1894. Arthroleptis hottegi Boulenger, Ann. Mus. Civ. Stor. Nat. Geneva,

(2) 15. p. 16, pi. iv, fig. 3: Auata Eiver, Somalia
Range. Ethiopia and Somalia.^*^^

Phrynobatrachus minutus (Boulenger)

1895. Arthroleptis minutus Boulenger, Proc. Zool. Soc. London, p. 539,

pi. XXX, fig. 4: "Durro," i.e. Duro, Ethiopia.
1910. Arthroleptis scheffleri Nieden, Sitzb. Ges. Naturf. Freunde Berlin,
p. 438: Nairobi and Kibwezi, Kenya Colony; Mpwapwa, Tan-
ganyika Territory; Zanzibar.
1924. Arthroleptis aliifer Ahl, Arch. Naturg., 90. Abt. A, p. 251:

Usaramo, Tanganyika Territory.
Range. Sudan; Somalia; *Uganda; *Kenya Colony; *Tan-
ganyika Territory ; Zanzibar Island ; *Northern Rhodesia ; Trans-
vaal (Mertens:1937) ; Angola (Monard:1937) ; *Belgian Congo.
Also recorded from French Equatorial Africa (Sternfeld :1917) ;
French Guiuea (Chabanaud :1921) and Portuguese Guinea
(Boulenger :1906), but all three require investigation.

Phrynobatrachus rxingwensis (Loveridge)

1932. Arthroleptis rwngwensis Loveridge, Bull. Mus. Comp. Zool., 72. p.
386: Ilolo, Eungwe Mountain, Tanganyika Territory.

Range. Southwestern *Tanganyika Territory and adjacent
*Belgian Congo.^^^

265 Records from Uganda (Peracca :1909 ; Boulenger :1911) and Belgian Congo
(Noble :1924) require verifying. I examined the Kibongoto, T. T. specimens of
Lonnberg (1907) and found they were based on minutus together with some
juvenile natalensis. The record from Singida, T. T. (FitzSimons :1930) was also
a juvenile natalensis.

266 A series of frogs from Lukafu received from the Congo Museum as gut-
terosus Chabanaud, a species described from French Guinea.

350 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Phrynobatrachus ukingensis mababiensis FitzSimons"®'

1932. Phrynobatrachiis mnbabicnsis FitzSimons, March, 1932, Ami.
Transvaal Mus., 15. p. 40, and 1935, ibid., 16, p. 390, fig. 26:
Tsotsoroga Pan, Mababe Flats, Bechuanaland Protectorate.
1940. Phryyiobatrachus vanrooyeni Hoffman, Soolog. Navors. Nas. Mus.
Bloemfontein, 1. p. 99, fig. 2B: Broedershoek, Greytown, Natal,
South Africa.
1944. Phrynobatrachus chiikilaensis Hoffman, Soolog. Navors. Nas. Mus.
Bloemfontein, 1. p. 177, figs. 5-6: "Chitiala," i.e. Chitala Elver,
Nyasaland.
Range. Marshes, chiefly upland, of southern *Tangan3'ika
Territory ; Mozambique ; *Nyasaland ; *Northern Rhodesia ;
*Beehuanaland ; *Natal ; *Belgian Congo.

Phrynobcrirachus uldngensis uldngenesis (Loveridge)

1932. Arthroleptis iikiitgaisis I.ovoridge, Bull. AIus. Comp. Zool., 72, p.

385: Madehani, Ukinga Mountains, Tanganyika Territory.
Range. Virgin forests of Ukinga and Ruugwe Mountains,
southern *Tanganyika Territory and Misuku Mountains, north-
ern Nyasaland.

Phxynobatrachus rouxi (Nieden)

1912. Arlliroleptis roiuri Nieden, Wiss. Ergebn. Deutsch. Zentral-Afrika-
Exped. 1907-1908, 4. p. 178, pi. v, figs. 5a-b: Northwest Buddu
Forest, Uganda.
Range. Uganda and adjacent *Belgian Congo.^^^

Genus ARTHROLEPTIS Smith ^"^

1849. Arthroleptis A. Smith, Illus. Zool. S. Africa, Kept., App., p. 24.

Type by monotypj': A. ica:hlbcrgii Smith.
1863. Hemimantis Peters, Monatsb. Akad. Wiss. Berlin, p. 451. Type by

monotypy: E. calcaratus Peters.

267 Frogs from N.vainkolo, southeast end of Lake Tanganyika, Northern Rho-
desia, topotypic of moorii Boulenger — to which I (1933) referred them in error
— are referable to mababiensis. The extensive series from the Uzungwe, Ubena
and Ukinga Mountains assigned by me (1933) to parrulus Boulenger, are also
mababiensis, the description of which appeared onl.v after my manuscript had
gone to press. Subsequently I had the opportunity of examining the types of
moorii and parrnlus in the British Museum.

268 The Mount Kenya record of Angel (1920) is considered doubtful.

269 Laurent (1954, Ann, Mus. Congo, Zool., 1. Misc. Zool. H. Schouteden, p.
3.'j) presents a chart setting forth the osteological characters hy which he sep-
arates t<choutedcnelht. at one end of the series, from Arthroleptis : his own (Coraeo-
diehus being intermediate between the extremes. His arrangi'ment involves the
transfer to Schoiitcdenclhi of at least a dozen Arthroleptis. The differences
(doubtless with nmiiy minor variants yet to be discovered) are of such a nature
that a subgeneric status appears to be the more lo^'ical way to escape the con-
fusion arising from frequent transfer of species from one "genus" to another.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 351

1921. Schoutedcnella Witte, Revue Zool. Afr., 9, p. 18. Type by mono-

typy: S. glohosa Witte.
1925. Pararthrolepti^ Ahl, Sitzb. Ges. Naturf. Freunde Berlin, 1923, p.

98. Type by present designation : P. nanus Ahl.
1939. Micrarthroleptis Deckert, Sitzb. Ges. Naturf. Freunde Berlin,

1938, p. 166. Type by original designation: Arthroleptis pyg-

maeus Alil.

1939. Pseudarthroleptis Deckert, Sitzb. Ges. Naturf. Freunde Berlin,

1938, p. 167. Type by original designation: "Arthroleptis,"
i.e. Hemimantis calcaratus Peters.

1940. Abroscaphm Laurent, Revue Zool. Bot. Afr., 34, p. 82. Type by

original designation: Arthroleptis adolfifrederici Nieden.
1940. Coracodichus Laurent, Revue Zool. Bot. Afr., 34. p. 85. Type by

original designation: Arthroleptis lohytli Boulenger.
1940. Arihroleptulus Laurent, Revue Zool. Bot. Afr., 34. p. 87. Type by

present designation: Arthroleptis xenodactylus Boulenger (also

included is the specimen of xenodaetyloides nkukae misidentified

as A. schuhotzi by Loveridge).

Arthroleptis xenodactylus Boulenger^'"

1909. Arthroleptis xenodactylus Boulenger, Ann. Mag. Nat. Hist., (8)

4. p. 496 : Amani, Usambara Mountains, Tanganyika Territory.

1910. Arthroleptis schubotzi Nieden, Sitzb. Ges. Naturf. Freunde Ber-

lin, p. 440 : Usumbura, Belgian Ruanda-Urundi.

1954. Schoutedcnella sylvatica Laurent, Ann. Mus. Congo, Zool., 1. Mis-
cellanea Zool. H. Sohouteden, p. 38: Buta, Uele, Belgian
Congo.

Range. Forest floor from sea level to 6000 feet in *Tanganyika
Territory ; Belgian Ruanda-Urundi and the *Belgian Congo.

Arthroleptis xenodaetyloides nkukae Loveridge

1942. Arthroleptis xenodaetyloides nkukae Loveridge, Bull. Mus. Comp.
Zool., 91, p. 427: Nkuka Forest, Rungwe Mountain, Tanganyika
Territory.
Range. Poorest floor in the Uzungwe ; Ukinga and Rungwe
Mountains, southern *Tanganyika Territory.

Arthroleptis reichei Nieden

1910. Arthroleptis reichei Nieden, Sitzb. Ges. Naturf. Freunde Berlin,
p. 437: Crater Lake, Ngosi Volcano, Poroto Mountains, Tan-
ganyika Territory.

Range. Virgin forests of Uzungwe ; Ukinga ; Rungwe and

270 I have compared topotypes of xenodactylus with topotyi>€3 (M.C.Z. 21718-
21) of sylvatica from the original series collected by de Witte (in February,
1925) and see no justification for regarding sylvatica as distinct.

352 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Poroto Mountains, southern * Tanganyika Territory and Misuku
Mountains, northern *Nyasaland.

Arthroleptis poecilonohis Peters

1863. Arthroleptis poecilonotus "Schlegel" Peters, Monatsb. Akad.

Wiss. Berlin, p. 446: Boutry, Ashanti, Gold Coast.
1882. Arthroleptis macrodactylus Boulenger, Cat. Batr. Sal. Brit. Mus.,

p. 117, pi. xi, fig. 5: Gabon, French Congo.
1885. Arthroleptis Mvittatus F. Miiller, Verb. Naturf. Ges. Basel, 7.

p. 671, pi. ix, figs, k-1: Tumbo Island, French Guinea.
1900. Arthroleptis inguinalis Boulenger, Proc. Zool. Soc. London, p.

442, pi. xxvii, fig. 2: Benito Eiver, Gabon, French Congo.
Range. Virgin forests of southern *Sudan ; *Uganda"'^? ; west
through *Belgian and *French Congo ; Fernando Po ; *French
and British Cameroon ; *Nigeria ; Dahomey ; Togo ; Gold Coast ;
*Ivory Coast; *Liberia; and *Sierra Leone to Portuguese
Guinea.

Artholeptis adoliifriderici adolfifriderici Nieden^'^

1910. Arthroleptis adolfi-friderici Nieden, Sitzb. Ges. Naturf. Freiinde

Berlin, p. 440: Eugege Forest, Belgian Euanda-Urundi.
1939. Arthroleptis affinis Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p. 303.
fig. 1: (Type ? 39 mm.) Amani, Usambara Mountains, Tan
ganyika Territory.
1939. Arthroleptis schocneheclci Ahl, Sitzb. Ges. Naturf. Freunde Berlin,
p. 305, fig. 2: (Type $ 22 mm.) Amani, Usambara Mountains,
Tanganyika Territory.
Range. Virgin forests of (Mbololo Mountain) *Kenya Colony
and (Usambara; Magrotto; Uluguru and Poroto Mountains)
*Tanganyika Territory ; west to Belgian Ruanda-Urundi and the
*Belgian Congo.

Arthroleptis stenodactylus whytii Boulenger-"

271 Included on the basis of a not-too-well-preserved frog (M.C.Z. 25402) that
I took at Mubango, Mabira Forest.

272 Trinomials are employed because of a southeastern race {A. a. jrancei
Loveritlge :1953) occurring in Ruo River forest, Mlanje Mountain, Nyasaland.

273 This montane-forest, burrow-breeding Arthroleptis differs from typical
s. stenodactylus in having a metatarsal tubercle that is usually shorter than,
though occasionally as long as, the inner toe ; also the tibio-tarsal articulation
of its forward-pressed hindlimb reaches the tympauiim or, rarely, the eye. On
the other hand, the lowland-savanna, burrow-breeding, s. stenodactylus has a
metatarsal tubercle that is usually longer than the inner toe, though in juveniles
it is occasionally only as long as the toe ; the tibio-tarsal articulation of its
adpressed hindlimb reaches only to the elbow or shoulder, very rarely as far as
the tympanum.

Owing to the difficulty of separating juveniles, and the general similarity of
the two forms, much confusion exists in the literature. A. whytii itself was a
composite of the two races ; ulugiiruensis was described by me owing to reliance
on Ahl's advice when he said that my Uluguru frogs were not subspccitically
identical with lonnhergi — of which I had no cotypes at that time. IS'ow the
Museum of Compai-ative Zoology has many topotypes of both whytii and all its
synonyms.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 353

1897. Arthroleptis whyiii Boiilcnger, Proc. Zool. Soe. London, pp. 801,
802, pi. xlvi, fig. 3: '"Masuka," i.e. Misuku Mountains, Nyasa-
land (restricted).
1915. Arthroleptis Jonnbergi Nieden, Mitt. Zool. Mus. Berlin, 7. p. 3G1 :

Mombo, foot of Usambara Mountains, Tanganyika Territory.
1932. Arthroleptis stenodactylus uluguruensis Loveridge, Proc. Biol. See.
Washington, 45, p. 61 : Nyingwa, Uluguru Mountains, Tan-
ganyika Territory.
1939. Arthroleptis vagus Ahl, Sitzb. Ges. Naturf. Freunde Berlin, p.
306: (Cotype $ 31 mm.; ^ 38 mm.) Usambara Mountains,
Tanganyika Territory.
1939. Arthroleptis iikamiensis Ahl, Sitzb. Ges. Naturf. Freunde Berlin,
p. 308, fig. 3: (Type $ 33 nun.) Ukami, i.e. Uluguru Mountains,
Tanganyika Territory.
Range. Mrgin forests of Usambara ; Magrotto ; Uluguru ; Ron-
do and Rungwe Mountains, *Tanganyika Territory ; also Misuku ;
Nchisi; Mlanje and Cliolo Mountains, *Nyasaland; Mozambique
and C'hirinda Mountain, *Southern Rhodesia.^"

Arthroleptis stenodactylus stenodactylus Pfeffer'''

1893. Arthroleptis stenodactylus Pfeffer, Jahrb. Hamburg. Wiss. Anst.,

1892, 10. p. 93, pi. i, fig. 2: Kihcngo, Tanganyika Territory.
192-1. Arthroleptis methneri Ahl, Arch. Naturg., 90, Abt. A, p. 251:
Xaugoma Cave, Matumbi near Kilwa, Tanganyika Territory.
1927. '*: Arthroleptis elegans Calabresi, Atti Soc. Ital. Sci. Xat. (Milano),

66, pp. 16, 35: Hongolo, Uadi Hoor, Somalia.
Range. Dry savanna, chiefly on the coastal plain, of *Kenya
Colony ; *Tanganyika Territory ; Mozambique ; *Nyasaland :
*Northern Rhodesia ; *Belgian Congo."^

Genus CACOSTERNUM Boulenger

1887. Cacosternum Boulenger, Ann. Mag. Nat. Hist., (5) 20, p. 51. Type
by monotj'py : C. namim Boulenger.

-"■•Credibly reported from the Belgian Congo by de Witte (iyo4), though
possibly typical stenodactylus of which we have examples from Kan.vama. B. C.

275 A., elegans, described from four 19 mm. frogs, is tentatively listed in the
synonymy in the hope that someone, bearing in mind the possibility they may
be juvenile s. stcnodtirtylus, will restudy the material.

276 Also reported from French Cameroon and Sierra Leone by Angel (1940),
possibly owing to confusion with raridbili.'i Matschie.

354 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Cacosternum boettgeri boettgeri (Boulenger)^"

1882. Arthroleptis ho.ettgeri Boulenger, Cat. Batr. Sal. Brit. Mus., p.

118, pi. xi, fig. 6: Vleis, Kaffraria, Bechuanaland Protectorate.

1887. Cacosternum nanum Boulenger, Ann. Mag. Nat. Hist., (5) 20, p.

52: Vleis, Kaffraria, Bechuanaland Protectorate.
1914. Arthroleptis schebeni Nieden, Sitzb. Ges. Naturf. Freunde Berlin,

1913, p. 438: Klein Nauas, Southwest Africa.
Range. Occurs in both upland savanna and arid regions of
East Africa, viz. Somalia ; *Uganda ; *Kenya Colony ; Bechuana-
land Protectorate ; *Transvaal ; Natal ( ? and/or the race albi-
venter Hewitt); *Orange River; Basutoland ; *Cape Province;
*Southwest Africa.

Genus HEMISUS Glinther

1858. Hemisus Giinther, Cat. Batr. Sal. Brit. Mus., p. 47. Type by

monotypy: Engystoma guttatum Rapp.
1863. Eakophrymis Steindachner, Sitzb. Akad. Wiss. Wien, 48, Abt. 1, p.

191. Type by monotypy: K. sudanesis (sic) Steindachner =

Engystoma nuirmoratum Peters.

Hemisus marmoratus marmorcrtus (Peters)
Eastern Sharp-snouted Frog
18.')4. Engystoma marmoratum Peters, Monatsb. Akad. Wiss. Berlin, p.

628 : Cabaeeira, Mozambique.
1863. Kakoplirynus sudanesis (sic) Steindachner, Sitzb. Akad. Wiss.

Wien, 48. Abt. 1, p. 191, pi. i, figs. 10-13: Sudan.
1882. Hemisus taitanus Peters, Reise nach Mossambique, 3. p. 175:

"Taita, " i.e. Teita, Kenya Colony.
Range. *Sudan ; Eritrea ; *Kenya Colony ; *Tanganyika Ter-
ritory ; Zanzibar Island ; *Mozambique ; *Nyasaland ; Southern
Rhodesia; Bechuanaland Protectorate; *Transvaal; southeast
Belgian Congo (Angola records are presumably referable to
the western race).

277 From Kundeluugru Plateau research station at 1750 metres, in Katanga,
southeast Belgian Congo. Laurent (1950, Revue Zool. Bot. Afr., 44, p. 138) has
described a single 14 mm. frog as C. lelcupL It is said to differ from boettgeri
by (1) lacking nuixillary teeth; (2) by the absence of markings on its hindlimbs.
It is true that limb markings are present in our extensive representation of
boettgeri from southern Africa. In East Africa, however, markings are both
present (M.C.Z. 2.5478 — 3 ex.) and absent (M.C.Z. 25479 — 2 ex.) in a series
from Kinangop Plateau (11,000 feet). In another Kinangop series markings are
present in 17 frogs, absent in 3 (M.C.Z. 16104, 16107, 10109). My colleague
Dr. E. E. Williams finds maxillary teeth present in all sizes of those he has
examined.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 355

Hemisus marmoratus guineensis Cope
Western Sharp-nosed Frog
1865. Hemisus guineensis Cope, Nat. Hist. Re\aew, 5, p. 100, footnote.

Type in Vienna Museum: Guinea (by inference).
1882. Ejigysloma vermiculatum "Schlegel" Peters, Reise nach Mossam-

bique, 3. p. 175: Ashanti, Gold Coast.
Range. * Western Tanganyika Territory; Angola; *Belgian
Congo ; French Congo ; French Equatorial Africa ; *French
Cameroon; Nigeria; Dahomey; *Gold Coast; *Liberia ; *Sierra
Leone ; French Guinea ; Portuguese Guinea ; Gambia ; Senegal.

Family PPIRYNOMERIDAE'^'
Genus PHRYNOMERUS Noble

1847. Brachymerus A. Smith (not of Dahlbom:1845), Illus. Zool. S.

Africa, Rept., text to pi. Ixiii. Type by monotypy: B. bifascia-

fus A. Smith.
1926. Phrynomerus Noble, Amer. Mus. Novit., No. 237, p. 20. New name

for Brachymerus A. Smith (preoccupied).

Phrynomerus bifasciatus bifasciatus (Smith)

1847.-"^ Brachymerus bifasciatus A. Smith, Illus. Zool. S. Africa, Rept.,

pi. Ixiii: "Country to the east and north-east of Cape Colony,"

i.e. east of Cape Province, South Africa.
1849.2^*^ Dendrobates inhambanensis Bianconi (1848), Nuovi Ann. Sci.

Nat., (2) 10, p. 107, ].l. V, litcs. 4-4a ; also 1850, Spec. Zool.

Mosamb., Rept., p. 26, pi. v. figs. 4-4a : Inhambane, Mozambique.
1944. Phrynomerus bifasciatus nyasalandensis Hoffman, Soolog. Navors.

Xas. Mus. Bloemfontein, 1. p. 181, fig. 9: "Chitiala," i.e.

Chitala River, Nyasaland.

-78 The status of the African genus Phrynomerus has been a chequered one.
In 1858 (as Brachymerus) it was made the type of a family (BRACHYMERI-
DAE) by Giinther (Cat. Batr. Sal. Brit. Mus., p. 124), onlv (as Phrynomantis)
to be merged in the family ENGYSTOMATIDAE by Boulenger (1882, Cat. Batr.
Sal. Ecaud. Brit. Mus., p. 172). In 19:51 (as Phrynomerus) it was given sub-
family rank (PHRYNOMERINAE) in the BRBVICIPITIDAE by Noble (Biologv
of the Amphibia, p. 538). later to be raised to full family status (PHRYNOMERI-
DAE) by Parker (1934: Monograph of the Frogs of the Family MICROHYLI-
DAE, p. 9) principally on account of its intercalary phalanges, a character it
shares with the RHACOPHORIDAE.

For other ways in which PHRYNOMERIDAE differs or agrees with MICRO-
HYLIDAE, together with a lucid tabulation of the characteristics of all African
firmisternal families, see Parker {loc. cit. pp. 8-9).

279 Actually appeared in December, 1847, fide Waterhouse (1880, Proc. Zool
Soc. London, p. 490).

280 Bianconi subsequently claimed that this description first appeared on
February 4, 1847. in Mem. Accad. Sei. 1st. Bologna, 1, p. 498 (a publication not
available to me).

356 BtTLLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Range. Kenya Colony; Tanganyika Territory; Zanzibar and
Mafia Islands ; south to Zululand ; west to Cape Province ;
northwest through Southwest Africa to Angola and the Belgian
Congo.

281

*o^

Family MICROHYLIDAE'"*'

Genus CALLULINA Nieden

1910. Callulina Nieden, Sitzb. Ges. Naturf. Freunde Berlin, p. 449. Type
by nionotypy: C. Jcreffti Nieden.

Callulina kreffti Nieden

1910. Calbilina kreffti Nieden, Sitzb. Ges. Naturf. Freunde Berlin, p.
449 : Amani, Usambara Mountains ; and Tanga, Tanganyika
Territory.

Range. Usambara, Magrotto and Uluguru Mountains, Tan-
ganyika Territory.

Genus SPELAEOPHRYNE A hi

1924. Spelacophryne Ahl, Zool. Anz., 61. p. 99. Type by original
designation: S. methneri Ahl.

Spelaeophryne methneri Ahl Scarlet-snouted Black Frog
1924. Spelaeophryne methneri Ahl, Zool. Anz., 61, p. 99: Nangoma

Cave, Matumbi near Kilwa, Tanganyika Territory.
Range. Eastern Tanganyika Territory.

Genus PROBREVICEPS Parker

1931. Prohreviccps Parker, Ann. Mag. Nat. Hist., (10) 8. p. 262. Type
by original designation: Breviceps macrodactylus Nieden.

Probreviceps macrodactylus macrodactylus (Nieden)

1926. Breviceps macrodactylus Nieden, Das Tierreich, 49, Anura, 2. p.

281 The western race microps Peters, has been reported from the Belgian
Congo, Sudan and the *Gold Coast (M.C.Z.).

282 This name, originally spelled MICRHYLIDAE by Giinther (1S5S, Cat. Batr.
Sal. Brit. Mus., p. 121), was amended and used by Parker (1943, Monograph of
the Frogs of the Family MICROHYLIDAE) for the family that, following Noble,
I have hitherto consistently called BREVICIPITIDAE of Cope (part). It is
typified by Rana gibbosa Linnaeus = Breviceps oibbosus (Linnaeus), the only
member of the family known to him in 1758.

The argument for retaining BREVICIPITIDAE advanced by Dunn (1935.
Copeia, p. 108) appears to be fallacious, being based on a misconception of
Parker's reasons for selecting MICROHYLIDAE. It is necessary only to add
that the sole species included by Giinther in his MICRHYLIDAE was Microhyla
achatina Tschudi of Java.

L0\T:RIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 357

G: Usambara Mountains, Tanganyika Territory.
1928. Breviceps usamharicu3 Barbour & Loveridge, Mem. Mus. Comp.

Zool., 50, p. 251 : Amani, Usambara Mountains, Tanganyika

Territory.
Range. Usambara Mountains, Tanganyika Territory.

Probreviceps macrodactylus loveridgei Parker

1931. Probreviceps macrodactylus loveridgei Parker, Ann. Mag. Nat.

Hist., (10) 8. p. 263: Bagilo, Uluguru Mountains, Tanganyika
Territory.

Range. Ulugnru Mountains, Tanganyika Territory.

Probreviceps macrodactylus rungwensis Loveridge

1932. Probreviceps macrodactylus rungwensis Loveridge, Bull. Mus.

Couip Zool., 72. p. 387: Nkuka Forest, Eungwe Mountain, Tan-
ganyika Territory.
Range. Rungwe Mountain, Tanganyika Territory.

Probreviceps uluguruensis (Loveridge)

1925. Breviceps iduguruensis Loveridge, Proc. Zool. Soc. London, p. 789,
pi. i, fig. 3 : Bagilo, Uluguru Mountains, Tanganyika Terri-
tory.
Range. Uluguru Mountains, Tanganyika Territory.

Genus BREVICEPS Merrem

1820. Breviceps Merrem, Vers. Syst. Amphib., p. 177. Type by mono-

typy: Pana gibbosa Linnaeus.
1826. Engystoma Fitzinger, Neue Class. Eept., pp. 39, 65. Type by

original designation: Sana gibbosa Linnaeus.
1830. Systoma Wagler, Nat. Syst. Amphib., p. 205. Type by monotypy:

Breviceps gibbosus Merrem = Sana gibbosa Linnaeus.

Breviceps mossambicus Peters Mozambique Short-headed Frog
1829. Engystoma grcunosum Cuvier (nomen nudum), Eegne Animal, ed. 2,

2, p. 112: "Cap," i.e. South Africa; probably Natal.
1854. Breviceps mossambicus Peters, Monatsb. Akad. Wiss. Berlin, p.

628: Mozambique Island, and Sena, Mozambique.
1944. Breviceps mitchelli Hoffman, Soolog. Navors. Nas. Mus. Bloem-

fonteiu 1, p. 182, fig. 10: "Chitiala," i.e. Chitala Eiver, Nyasa-

land.
Range. Tanganyika Territory, south to Natal, northwest
through Southern Rhodesia to Angola and the Belgian Congo.

358 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Genus PARHOPLOPHRYNE Barbour & Loveridge

1928. Parlioplopliryne Barbour & Loveridge, Mem. Mus. Comp, Zool.,
50. p. 260. Type by original designation: P. vsambaricus (sic)
Barbour & Loveridge.

Parhoplophryne usamborica Barbour & Loveridge

1928. Parhoplophryne usambaricus Barbour & Loveridge, Mem. Mus.
Comp. Zool., 50, p. 260: Amani, Usambara Mountains, Tan-
ganyika Territory.

Range. Usambara Mountains, Tanganyika Territory.

Genus HOPLOPHRYNE Barbour & Loveridge

1928. Hoplophryne Barbour & Loveridge, Mem. Mus. Comp. Zool., 50,
p. 253. Type by original designation : H. %dugyru.ensis Barbour
& Loveridge.

Hoplophryne rogersi Barbour & Loveridge
Usambara Banana-Frog
1928. Hoplophryne rogersi Barbour & Loveridge, Mem. Mus. Comp. Zool.,
50, p. 258, pi. ii, fig. 5 : Mount Bomoli, Amani, Usambara
Mountains, Tanganyika Territory.
Range. Usambara and Magrotto Mountains, Tanganyika Ter-
ritory.

Hoplophryne uluguruensis Barbour & Loveridge
Uluguru Banana-Prog
1928. Hoplophryne uluguruensis Barbour & Loveridge, Mem. Mus. Comp.
Zool., 50, p. 254, pi. ii, figs. 3-4: Mount Mbova, Nyange, Ulu-
guru Mountains, Tanganyika Territory.
Range. Uluguru Mountains, Tanganyika Territorj'.

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

359

LIST OF SPECIES OR RACES STILL UNREPRESENTED
IN THE MUSEUM OF COMPARATIVE ZOOLOGY,

CAMBRIDGE

LIZARDS

*Hemidactylus alhopunctaUis Loveridge
Ebennvia sp.

Chamaeleo hitae7u<ittis schuhoizi Stenifeld
Chamaelco fisdieri ntJimoUeri L. Miiller
Chamaeleo fischeri fischeri Beichenow
Mabuya varia brauni Tornier
Lygosoma Tcutuensis Lonnberg

*Philochortus intermedins rudolferisis Parker
Ichnotropis tamganicana Boulenger

SNAKES

*Leptotyphlops fiechteri Scortecci

^Telescopus dhara somalicus (Parker)
CrotaphopeJtis wrr7icri Boulenger

Calamelaps unicoJor polylepis Bocage

TOADS AND FROGS

Haud, British Somaliland
Chokocho, Pemba Island
Mt. Kenya at 14,000 feet
Mt. Hanang at 2,300 feet
Mt. Nguru, Tang. Terr.
Ukinga Mtns., Tang. Terr.
Kutu Village nr Fort Hall,

K.C.
Kaliokwell River, K.C.
Lake Tanganyika

Duke of Abruzzi 's Villa,

Somalia
Xgatana, Tana River, K.C.
Usambara Mountains, Tang.

Terr.
Tukuyu, s.w. Tang. Terr.

"Bii/o regularis mara]:iietensis Roux

Bufo lindneri Mertens
*Leptopell'< gramineus (Boulenger)
Hylambates verrucosus Boulenger

EyperoUu-s albifrons Ahl

HyperoUus alhofrenatus Ahl

Hyperolius albolabris Ahl

HyperoUus bergeri Ahl

Hyperolius glandicolor Peters

Hyperolius guttolineatu-s Ahl

Hyperolius marmoratus campylogrammus

*Hyperolius pachydermus Werner

Mt. Marakwet at 2500 m.,

K.C.
Dar es Salaam, Tang. Terr.
Badditu to Dime, Somalia
Mabira Forest, Kyagwe,

Uganda
"Africa" Mafia Island, T.T.
Tanganyika Territory
Kwa Buosch or Bnorch, T.T.
Uaso Narok, Kenya Colony
Teita, Kenya Colony
Tanganyika Territory
Laurent

Bura, 1050 m.,

Teita, K.C.
Gondokoro, Sudan ; also U. ;

K.C.

:j60 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Hyper olius pantherinus (Steinclachner) Laikipia, Kenya Colony

Eyperolius quadratomaculatus Ahl Molioro, Tanganyika

Territory

Eyperolius acriptus Ahl Tanga, Tanganyika Territory

Eyperolius tornicri Ahl Uluguru Mountains, T.T.

Eyperolius viridiflavus pitmani Laurent Lake Bunyoni, s.w. Uganda

Phrynobatrachus bottegi (Boulenger) Auata Eiver, Somalia

Phrynobatrachus rouxi Nieden N.W. Buddu Forest, Uganda

ANNOTATED BIBLIOGRAPHY

of revisionary studies or other contributions likel}^ to be useful
to anyone taking- up the study of East African herpetology.
Those selected have all been published since G. A. Boulenger 's
nine-volume series of catalogues on the collections in the British
Museum (1882-1896).

Boulenger, G. A.

1915. A List of the Snakes of East Africa, north of the Zambesi and
south of the Soudan and Somaliland, and of Nyassaland. Proc.
Zool. Soe. London, pp. 611-640, figs. 1-3. (Still the handiest
aid to an approximate identification of most snakes, though
the nomenclature is even more obsolete than the spelling of
Zambezi, Sudan and Nyasaland.)
1920- Monograph of the Lacertidae. (British Museum), 1. pp. x -f
1921. 352; 2, pp. viii -(- 451. (Contains good descriptions of most of
the 21 East African forms, but the nomenclature is binomial;
geographical races are treated either as full species or "varie-
ties," a term variously employed.)

Laurent, Eaymond

1950. Eevision du genre Atractaspis A. Smith. Mem. Inst. Eoy. Sci.
Nat. Belgique, (2) 38. pp. 1-49, figs. 1-18. (This well-illustrated
treatise subdivides somewhat excessively; see footnotes to pp.
296-297 of this check list regarding the East African forms.)
See also G. F. de Witte and E. Laurent.

LovERiDGE, Arthur

1923. A List of the Lizards of British Territories in East Africa . . .
with Keys for the Diagnosis of the Species. Proc. Zool. Soc.
London, pp. 841-863. (Moderately serviceable where not re-
placed by family revisions, as listed below.)

*An asterisk indicates that it occurs in the Northern Frontier Province of
Kenya Colony.

The Museum of Comparative Zoology would be glad to offer an exchange for
examples of any of the foregoing 32 desiderata.

LON^ERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS 361

1924. Check List of the Reptilia recorded from the British Territories
in East Africa. Journ. E. Africa Uganda Nat. Hist. Soc. Spec.
Suppl., No. 3, pp. 1-16. (Superseded.)

1930. A List of the Amphibia of the British Territories in East
Africa . . . with Keys for the Diagnosis of the Species. Proc.
Zool. Soc. London, pp. 7-32. (Rather useless; as at least 40
species have been added, and some removed, since 1930.)

1939. Revision of the African Snakes of the Genera Mehelya and
Gonionotophis. Bull. Mus. Comp. Zool., 86. pp. 131-162, figs. 1-2.
(Substantially up to date; some range extensions.)

1940. Revision of the African Snakes of the Genera Dromophis and
Psammophis. Bull. Mus. Comp. Zool., 87. pp. 1-70. (No change
as regards East African forms, but some workers disapprove
of scholcari and certain southwestern snakes being treated as
races of sibilans.')

1941. Revision of the African Lizards of the Family Amphisbaenidae.
Bull. Mus. Comp. Zool., 88. pp. 467-524, figs. 1-53. (See footnote
to AmphishaeTia in this check list.)

1941. Revision of the African Terrapin of the Family Pelomedusidae.
Bull. Mus. Comp. Zool., 88. pp. 467-524. (The West African
Pelusios niger should be removed from the synonymy of sub-
niger.)

1942. Revision of the Afro-Oriental Geckos of the Genus Phelswma.
BuU. Mus. Comp. Zool., 89. pp. 437-482, figs. 1-2, map.

1942. Revision of the African Lizards of the Family Gerrhosauridae.
Bull. Mus. Comp. Zool., 89. pp. 483-543. (Now merged with the
Cordylidae.)

1944. Revision of the African Lizards of the Family Cordylidae. Bull.
Mus. Comp. Zool., 95, pp. 1-118, pis. i-xii. (See also Gerrhosauri-
dae above.)

1944. Further Revisions of African Snake Genera. Bull. Mus. Comp.
Zool., 95. pp. 119-247. (Deals with Rhamnophis ; Thrasops;
Duherria; Thelotornis; Calamelaps ; Miodon; Aparallactus;
Elapsaidea and Paranaja.)

1947. Revision of the African Lizards of the Family Gekkonidae.
Bull. Mus. Comp. Zool., 98. pp. 1-469, pis. i-vii. (This revision,
like its predecessors, contains keys to the species together with
a synopsis of all that is known of their life history, enemies,
parasites, etc.)

LovEEiDGE, A., and E. E. Williams

1957. Revision of the African Tortoises and Turtles of the Suborder
Cryptodira. Bull. Mus. Comp. Zool., 115, pp. 163-557, figs. 1-62,
pis. i-xviii. (Together with the revision of the Pelomedusidae,
Suborder Pleurodira, this covers all the African Testudinata.)

362 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Mertbns, Robert

1942. Die Familie der Warane (Varanidae), III. Taxonomie. Abhand.
Senckenberg. Naturf. Ges., No. 466, pp. 237-391. (Nos. 462 and
465, the first two parts of this exhaustive treatise by a foremost
herpetologist, deal with the skull, etc. etc. They also appeared
in 1942.)

Mertens, Robert and Heinz Wermuth

1955. Die rezenten Schildkroten, Krokodile und Briickenechsen. Zool.
Jahrb., Syst., 83. pp. 323-440. (A check list of the turtles, tor-
toises, crocodiles and tuatara of the world.)

MORBAU. R. E. and R. H. W. Pakenham

1941. The Land Vertebrates of Pemba, Zanzibar, and Mafia: a Zoo-
geographical Study. Proc. Zool. Soe. London for 1940, 110
(A),pp. 97-128, fig. 1.

1942. Corrigenda and Addenda to "The Land Vetebrates of Pemba,
Zanzibar, and Mafia: a Zoogeographical Study." Proc. Zool.
Soc. London, 112. (A), pp. 61-63.

Paken-ham, R. H. W., see R. E. Moreau and R. H. W. Pakenham

Parker, II. W.

1934. A Monograph of the Frogs of the Family Microhylidae. London,
pp. viii -f 208, figs. 1-67.

Pitman, C. R. S.

1938. A Guide to the Snakes of Uganda. (Kampala), pp. xxi -|- 362,
pis. i-xxviii, col. pis. A-Q, diagrams I-II, maps. (The excellent
figures and coloured plates make this work invaluable to begin-
ners. A new and revised edition is in course of preparation as
the 1938 edition is out of print.)

Williams, Ernest E., see A. Loveridge and E. E. Williams

WiTTE, G. F. de, and R. Laurent

1943. Contriliution a la Systematique des Formes degrades de la
Familie des Scincidae apparentees au Genre Scelotrs Fitzinger.
Mem. Mus. Roy. Hist. Nat. Belgique, (2) 20. pp. 1-44, figs. 1-61.

1947. Revision d'un Groupe de Colubridae Africains (Genres Calame-
laps, Miodon, Aparallactus et Formes affines). Mem. Mus. Roy.
Hist. Nat. Belgique, (2) 29. pp. 1-134, figs. 1-132. (The conclu-
sions arrived at by these authors are often at variance with
those in my 1944 paper dealing with the same genera. A.L.)

INDEX

To reduce the bulk of this index, specific or racial names have
not been listed under the heading of the genera to which they
belong but only individually as species or races. To find all the
"East African" species represented in any given genus it will
be necessary to turn to the genus in question in the text.

abbotti, Chamaeleo, 204
aberae, Ptychadena, 341
aberdariensis, Rana, 339
Ablabophis, 249
Ablepharus, 218
Abrana, 338
Abroscaphus, 351
abyssiniea, Duberria I., 266
abyssinica, Eana, 342
abyssiuicum, Homalosoma, 266
abyssinicum, Lyeophidium, 252
abyssinus, Rachiodou, 288
acanthias, Urobelus, 283
Acanthixalus, 321
Acanthocalyx, 258
.\eanthocercus, 193
aeeiaeensis, Hylarthroleptis, 346
aehatina, Microhyla, 356
Aoontias, 222
aeontias, Melanoseps, 222
acontias, Scoleooseps, 222
acridoides, Phrynobatrachus, 346,

347
acridoides, Staurois, 346
Airidophaga, 300
aciileata, Agania, 158, 192
at'uleatus, Gecko, 185
aculeatus, Holodactylus, 179
acuticeps, Hyperolius, 333
acutirostre, Lycophidion c, 253
aeutus, Crocodilus, 176
acutus, Geocalamus, 239
aeutus, Onychocephalus, 240
aeutus, Psamniophis, 277
acutus, Rhamphiophis a., 277
Adenomus, 309
adolfi, Dicroglossus, 337

adolfi, Typhlops, 240, 242
adolfifriderici, Arthroleptis, 351, 352
adolfif riderici, Arthroleptis f ., 352
Adolfus, 228
adspersa, Rana a., 344
adspersus, Pyxicephalus, 336
aegyptiacus, Trionyx, 171, 172
aegyptius, Trapelus, 193
Aelurophis, 270
aemulans, Cynodontophis, 283
aenea, Cricochalcis, 226
aesculapii. Coluber, 257
aesculapii, Zamenis, 257
aethiopissa, Rliamnophis, 264
afer, Eumeces, 215
affinis, Arthroleptis, 352
affinis, Chamaeleon j., 205
affinis, Hemidactylus, 184
afghanus, Polypedates, 337
afrieana, Agama agania, 194
afrieana, Caitia, 225
afrieana, Cnemaspis a., 180
afrieana, Fordia, 171, 172
africanus, Ablepharus b., 218, 210
africanus. Algiroides, 228
africanus, Algyroides, 228
africanus, Crocodylus, 177
africanus, Gymnodaetylus, 180
africanus, Holodactylus, 178
Afrixalus, 321, 322, 324
agadezi, Arthroleptis, 345
Agalmatosaurus, 235
Agama, 192
agama, Agama a., 194
agama, Lacerta, 192, 194
AGAMIDAE, 192, 196
agassizii, Chelonia, 16fi

11

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Agetosaura, 229

agilig, Agama, 192

agilis, Lacerta, 229

agilis, Scincus, 208

AGLOSSA, 307

agua, Bufo, 309

ahli, Arthroleptis, 345

ahli, Hyperolius, 331

ahli, Hyperolius a., 331

Ailurophis, 270

albescens, Chiromantis, 316

albifer, Arthroleptis, 349

albifrons, Hyperolius, 334, 359

albifrons, Stenostoma, 245, 246

albigularis, Tupinambis, 235

albigularis, Yaranus e., 236

albofrenatus, Hyperolius, 334, 359

albolabris, Hyperolius, 334, 359

albolabris, Eana, 338

albolabris, Kana a., 338

albopunctatus, Euprepes o., 212

albopunctatus, Hemidactylus, 182,

359
albopunctatus, Scaphiophis, 264
albopunctatus, Scaphiophis a., 264
albotaeniata, Mabuia, 210
:xlbotaeniata, Mabuya m., 210
albovariata, Dendrophis, 261
Algiroides, 228
Algyroides, 228
Aliurus, 182
alleni, Algiroides, 228
alleni, Algyroides, 228
alleni, Pseudoxenopus, 338
alleni, Eiina c, 338
alluaudi, Hemidactylus, 183
Alopeeion, 250

aloysii-sabaudiae, Lygosoma, 216
Alsodromus, 229
altaeelgouis, Chamaeleo b., 201
altaeelgoiiis, Chamaeleon b., 201
attarensis, Leptodira, 272
alteruans, Hydrophis b., 295
alticola, llyjieroliiis, 326

amaniensis, Leptopelis, 319
Amblyodipsas, 282
americana, Pentonix, 174
americanus, Crocodilus, 176
amethistina. Boa, 247
ammodytes. Coluber, 300
Amolops, 337
AMPHIBIA, 305
Amphiglossus, 220
Amphiophis, 279
Amphisbaena, 237
AMPHISBAENIDAE, 237
Amphisbaenula, 238
Amplorliinus, 275
Analcis, 288
ANAPSIDA, 163
Anaxyrus, 309
anchietae, Chamaeleo, 198
anchietae, Chamaeleo s., 198
anchietae, Eumecia, 213, 214, 216
anchietae, Euprepes, 209
anchietae, Hylambates, 317
anchietae, Leptopelis, 317
anchietae, Naja h., 292
anchietae, Ophirhina, 266
anchietae, Rana, 340
anchietae, Riopa, 216
Ancylocranium, 237
andersonii, Atractaspis, 296
Anelytrops, 222
angasijanus, Euprepes, 210
angeli, Atractaspis i., 297
angeli, Cassina, 321
angolensis, Amphiophis, 279, 280
angolensis, Causus r., 299
angolensis, Euprepes, 212
angolensis, Hylambates, 317
angolensis, Lygodactylus, 187
angolensis, Onychocephalus, 242
angolensis, Philothamnus, 261
angolensis, Psammophis, 280
angolensis, Rana, 339
angolensis, Rana f., 339, 340
angolensis, Sepsina, 220

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

111

anguina, Lacerta, 226
anguineus, Bipes, 210
anguineus, Herpetoseps, 220
anguiniformis, Boa, 248
angularis, Lygodactylus, 188
jingularis, Lygodactylus a., 188
;uigulatus, Hemidaetylus, 184
angulatus, Hemidaetylus b., 184
augulifera, Dinodipsas, 299
angusta, Sphargis, 164
aiigusticeps, Dendroaspis, 294
angustieeps, Naia, 294
angustieoronatus, Chamaeleo, 200
augustirostris, Dicroglossus, 348
Anilios, 240
anisolepis, Atheris, 304
annectens, Agama, 195
aimeetens, Chamaesaura, 226
annectens, Lygodactylus s., 186
aunulifer, Alopecion, 250
annulifera, Naja h., 291
Anodon, 288
anomalus, Bufo, 310
anomalus, Morethia, 218
Aiioplodipsas, 269
Anoplopus, 190
anotis, Bufo, 313
Ansonia, 310
ansorgii, Rana, 343
antinorii, Dendraspis, 294
antinorii, Denroaspis p., 294
Apai-allactus, 268, 284
Apathya, 231
Apistes, 228
APODA, 305
Apola, 196
Aporarchus, 237
aquaticus, Bufo, 308
arabicus, Boaedou f ., 251
Archaius, 197
ARCHOSAURIA, 176
arctiventris. Coluber, 266
areuata, Testudo, 164
arenicola, Echis. 303

arenicola, Herpetosaura, 220
arenicola, Typhlops b., 244
argenteus, Hylambates, 317
argenteus, Leptopelis, 317
argentii, Nubilia, 181
aigcntovittis, Hyperolius, 330
ai-gus, Hyperolius, 331
argus, Hyperolius a., 331
arguta, Lacerta, 233
argyreivittis, Cystignathus, 321
Argyrogena, 257
Argyrophis, 240
arietans, Bitis a., 159, 273, 301
arietans. Echidna, 301
arietans, Vipera, 302
armata, Agama, 193
armata, Agama h., 193
armatus, Gekko, 185
Arthroleptides, 335
Arthroleptis, 214, 335, 350
Arthroleptulus, 351
asper, Bufo, 309
asper, Hyperolius, 329
Aspidoboa, 248
Aspidochelys, 172
Aspidonectes, 171
Aspidopython, 247
Aspidorhinus, 232
Aspidorhj-nclius, 239
aspilus, Aspidonectes, 172
Aspis, 291
aspis. Coluber, 300
assimilis, Dipsadoboa, 269
Asterophis, 248
Asthenophis, 265
astrolabi, Amphiglossus, 220
Astylosternus, 214
ATELOPODIDAE, 308
ater, Typhlops, 239
aterrima, Atractaspis, 297
ATHECAE, 163
Atheris, 159, 303
altantica, Dermochelis, 164
atlantieus, Scoliophis, 257

IV

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

atia, Agconia, 192
.'lira, Caret ta, 167
atra, Dasypeltis s., 288
atra, Herpetosaura, 221
Atractaspis, 296
atratus, Neusterophis, 256
atrieeps, Naja ii., 292
atricollis, Agama, 195
atriventris, Homalosoma 1., 266
Atropis, 229
atropos, Coluber, 301
attarensis, Leptodira, 272
attenuatus, Seoleeomorphus, 396
Aubria, 337
aubryi, Cryptopus, 172
aubryi, Hyla, 316, 318
aubryi, LfCptopelis, 31 S
audax, Hormonotus, 253
aulicus, Chaniaetortus, 272
aulicus, Chaniaetortus a., 272
auritus, Coluber, 279
auritus, Phryuoliatracluis, 347
australis, Gerrliosaurus n.. 225
australis, Kinixys, 170
babaulti, Atrai-tasi)i.s, 297
babeoc-ki, Testudo p., 169
badia, Testudo, 173
badioflavus, Adeuonuis, 309
bahiensis, Natrix, 257
Baliopygus, 337
barbouri, Atheris, 305
barbouri, Hemidaetylus s., IS-J
barbouri, Hemidaetylus t., 183
barbouri, Leptopelis, 318
barbouri, Eaua, 341
barkeri, Ancyloeraniuiu, 237
barnumbrowni, Tarbophis, 272
barotseensis, Naja h., 292
barrowii, Onychopliis, 242
Barycephalus, 193
Baseanion, 257
Baseanium, 257
batesii, Aparallactus, 286
l)ntilliferus, Uromastix, 193

battersbyi, Philothamnus i., 260
bayoni, Hyperolius, 326
bayoni, Rappia, 326
bayonii, Dumerilia, 220
bayonii, Hemidaetylus, 184
bayonii, Mabuya b., 211
Bdellophis, 306
beeearii, Paleorana, 338
bechuanieus, Pelusios, 175
Bedriagaia, 227
Ijelliana, Kinixys, 170
belliana, Kinixys b., 170
liellii, Bueephalus, 273
bellii, Herpetaethiops, 260
l)eng;ilensis, Tupinambis, 235
In'iiguelensis, Bufo, 311
beiiguellensis, Hemidaetylus, 185
bcunettii, Mecistops, 177
bequaerti, Cliauiaeleon, 201
bequaerti, Chlorophis, 262
bergeri, Chamaeleon, 201
t)ergeri, Hyperolius, 334, 359
bergeri, Mabuia m., 210
bergeri, Pseudoprosymna, 265
bergi, Gerrhosaurus, 224
bergii, Paehydactylus, 191
Berus, 300
berus, Coluber, 300
Bettaia, 230

bianconii, Dasypeltis s., 289
bibronii, Atractaspis, 296, 298
liilironii, Atractaspis 1j., 298
bicariuata, Chelouia., 165
Ijicarinatus, Heterolepis, 255
bicolor, Argyrophis, 240
!)icolor, Coluber, 272
liicolor, Glypliolycus, 250
lucolor, lleteururus, 269
bicolor, Hydrus, 295
lucolor, Lycodonomorphus, 250
bifolor, Pelamis, 295
Ificoloratus, Micrelaps, 282
lU 'uspis, 197
biiascintus, Bi'ai'hymerus, 355

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

bifasciatus, Phrynonierus b., 355
hifidus, Chamaeleon, 196, 197
bifurcatus, Diecros, 196
bifurcus, Clianiaeleo, 197
bilineatus, Causus r., 299
biliueatus, Psammophis in., 280
liiliueatus, Typhlops, 245, 246
bilobus, Clianiaeleo, 200
binuensis, Crocodilus, 177
bipes, Anguis, 219
bipostocularis, Atractaspis, 297
bipostocularis, Atractaspis i., 297
bipraeoeularis, Boodon, 251
biscutatus, Crocodilus, 176
biseriatus, Psammophis, 280
biseriatus, Psammophis b., 280
liissa, Caretta, 166
bitaeniatus, Cluimaeleo, 200
bitaeniatus, Clianiaeleo b., 200
bitaeniatus, Ilyperolius, 328
bitaeniatus, Psammophis, 280
bitin. Coluber, 301
Bitis, 301
bitis, Coluber, 301
bitorquata, Coronella, 259
bivittata, Ichiiotropis, 234
bivittata, Ichnotropis c, 234
bivittatus, Arthroleptis, 352
bivittatus, Chamaeleo, 200
bivittatus, Ilyperolius, 322
bivittatus, Sciucus, 218
bivittatus, Tupiuambis, 235
blandingii, Boiga, 268
blandingii, Uipsas, 268
l)lanfordii, Typhlops, 240
blanfordii, Typhlops b., 240
Blepharosteres, 218
Boaedon, 250

bocagei, Hemidactylus, ISl!
bocagei, Typhlops, 242
boeagii, AhaetuUa, 262
bocagii, Cystignatlius, 316, 317
boeagii, Leptopdis, 317
bocagii, .Mal)uya, 212

bocagii, Prosymna, 265
bocagii, Prosymna a., 265
bocagii, Psammophis, 280
boettgeri, Arthroleptis, 354
boettgeri, Cacosteruum b., 354
boettgeri, Rhampholeon, 207
bottgeri, Rina t., 341
bohmanni, Cnemaspis, 180
BOIDAE, 247
boiei, Xenopus, 307
Boiga, 159, 268, 271
BOIGINAE, 268
BOINAE, 248
tiojeri, Hemidactylus, 183
i)0,ierii, Scincus, 220
Boltalia, 181
Bombifrons, 176
bombifrons, Crocodilus, 176
bonebfrgi, Natalobatrachus, 346
liorbonica, Ilyla, 308
borealis, Xenopus 1., 307
liosniensis, Vipera b., 300
Bothrophthalmus, 249
bottegi, Arthroleptis, 349
bottegi, Phrynobatrachus, 349, 359
bottegi, St^rnothaerus, 175
liottegoi, Gerrhosaurus, 224
bottegoi, Gerrhosaurus m., 224
boueti, Dipsadomorphus, 269
boulengeri, Algiroides, 229
boulengeri, Aparallactus, 285
boulengeri, Aparallactus c., 286
boulengeri, Boulengerula, 305, 306
boulengeri, Brookesia s., 207
Ijoulengeri, Cnemaspis, 180
boulengeri, t^lapsoidea, 290
boulengeri, Feylinia, 222
boulengeri, Glauconia, 247
boulengeri, Leptotyphlops, 246
boulengeri, Mabuia, 210
boulengeri, Mabuya, 210 •
boulengeri, Nucras, 230
boulengeri, Nucras b., 230
boulengeri, Pachydactylus, 19^'

VI

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

boulengeri, Phrynobatraehus, 346
})Oulengeri, Rhampholeon, 207
boulengeri, Scolecoseps, 222
boulengeri, Typhlops, 242
Boulengeria, 232
boulengerii, Phrynopsis, 337, 344
Boulengerina, 290
Boulengerula, 305
bowringii, Doryura, 181
Brachyblemma, 266
brachycnemis, Afrixalus b., 322, 323
brachycnemis, Megalixalus, 323
Brachycranion, 296
BRACHYAIERIDAE, 355
Brachymerus, 355
Brachysaura, 193
brachyura, Vipera, 302
liradfieldi, Lygodaetylus, 187
Bradypodion, 196, 197
Bradj-podium, 196, 197
bragantina, Eana, 344
liraminus, Eryx, 244
hraminus, Typlilops, 244
lirasiliana, Bionia, 237
braueri, Glauconia, 245
brauni, Bufo, 310
brauni, Mabuia, 212
brauni, Mabuya v., 212, 359
bravana, Eana g., 338
bravanus, Limnodytes, 338
bredoi, Latastia, 231
brevicaudata, Brookesia, 207
brevicaudatus, Chamaeleon, 207
Breviceps, 357
breviceps, Dasypeltis, 288
breviceps, Hj'perolius, 329
breviceps, Raua, 337
BREVICIPITIDAE, 355, 356
brevicollis, Euprepes, 211
brevieollis, Mabuya, 211
brevipalmatus, Hylambates, 317
brevipahnatus, Hyperolius, 323
brevirostris, Chanipse, 176
brevirostris, Psammophis, 276, 279

brevis, Typhlops, 241

brevis, Typhlops s., 240

brieni, Hyperolius, 330

Bronia, 237

Brookesia, 206

brunnea, Pelamis p., 296

Bucephalus, 273

budduensis, Leptopelis, 318

Bufo, 308, 309, 310

bufo, Rana, 309

bufonia, Maltzania, 337

BUFONIDAE, 308

Bunocnemis, 182

bunyoniensis, Xenopus 1., 308

burchelli, Chamaeleo, 198

burchelli, Eremias, 232

burgeoni, Chamaeleon, 201

burgeoni, Lygosoma, 217

burgeoni, Rappia, 325

burtonii, PoeciJostolus, 303, 304

burtoni, Tiliqua, 211

butleri, Chilorhinophis, 284

butleri, SimocephaJus, 254

Cacophryne, 308

Cacosternum, 163, 353

Cadmus, 266

caeca, Amphisbaena, 237

CAECILIIDAE, 305

eaeruleopunctatus, Hyperolius, 329

caerulescens, Coluber, 261

caerulescens, OUotis, 310

caeruleus, Coluber, 201

caeruleus, Hapsidophrys, 263

caesar, Xenurophis, 267

caesius. Coluber, 261

Caetia, 225

caeutiens, Elapomorphus, 283

caffer, Sepomoi'phus, 220

Caita, 225

calabaricus, Heniidactylus f., 185

Calamelaps, 281

calamita, Bufo, 309

ealcaratus, Arthroleptis, 350

calearatus, Hemimantis, 350, 351

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

Vll

Calechidna, 301

callichromus, Hyperolius, 330

Calhiliiia, 35G

Caliiinnia, 197

f-alyptratus, Chamaeleo, 197

CaIjT)trosaura, 197

Camaeleo, 196

Camaeleon, 196

Canieleo, 196

camerunensis, Bufo c, 311

camerunt'iisis, Meizodon c, 2o9

Campsodactylus, 213

campylogranunus, Ilyperolius ni.,

335, 359
eana, Pseudaspis, 266
candidissinuis, Coluber, 291
eaudidus, Cerastes, 275, 291
Cantinia, 191
canus, Coluber, 266
Coauana, 167
caouana, Testudo, 167
caouana, Tlialassochelys, 167
capellii, Chamaeleo, 200
capense, Lycophidion c, 252, 253
capensis, Algyra, 233, 234
capensis, Aparallaetus, 284, 285, 287
capensis, Aparallaetus c., 287
capensis, Bucephalus, 273
capensis, Dasji^eltis s., 288
capensis, Elapomorphus, 285, 287
capensis, Gongylus, 220
capensis, Hemidactylus, 187
capensis, Herinia, 208, 220
capensis, Heterolepis, 254
capensis, Laeerta, 232, 236
capensis, Lycodon, 251, 252
capensis, Lygodactylus, 187
capensis, Mabuya, 220
capensis, Mehelya c, 254, 255
capensis, Onychocephalus, 244
capensis, Pentonyx, 173
capensis, Phrynobatrachus, 346
capensis, Stellio, 196
capensis, Tarcntola, 191

capensis, Thelotornis, 274
capensis, Thelotornis k., 274
capensis, Tiliqua, 208, 220
capensis, Tropidosaura, 233
capensis, Typhlops, 245
capistratus. Coluber, 257
eappadocia, Laeerta, 231
caracal, Gecko, 183
carens, Bufo, 309, 310
Caretta, 166, 167
caretta, Caretta, 167
caretta, Testudo, 167
carinata, Pseudoboa, 303
carinatus, Chlorophis, 260
carinatus, Dipsas, 289
carinatus, Echis c, 303
carinatus, Pliilothamnus h., 260
carinatus, Scincus, 208
cariniventris, Agama, 194
carpoutcri, Chamaeleo, 203
carsonii, Ablephaius, 219
caspius. Coluber, 257
Cassina, 320
Cassiuiopsis, 321
castanea, Emys, 174, 175
castiinea, Kinixys, 170, 171
castanoidcs, Pelusios n., 175
Catapherodon, 250
cataphraetus, Crocodilus, 176, 177
cataphractus, Crocodylus, 176
Cathetorhinus, 240
Catodon, 246
eaudospina, Agama, 195
caudospiua, Agama p., 19."J
Causus, 298
Centrochelys, 168, 169
Centromastix, 229
cepedinna, Testudo, 165
cepediauus, Gecko, 190
Cephaloehelys, 167
Cephalophis, 268
cephalo, Testudo, 167
cephalo, Thalassochelys, 167
Ceramodactylus, 179

Vlll

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Cerastes, 159, 275

c-eratophorus, Atheris, 305

Cercocalamus, 285

eeylanicus, Hoplobatraclms, 337

Chal)iinaudia, 222

Chaelio, 196

Chamaeleo, 158, 196, 197

Cliainaeleon, 190

CHAMAELEONIDAE, 196

chamaeleon, Lacerta, 196

CHAMAELEONTES, 196

CHAMAELEONTIDAE, 196

Ohaniaesauia, 226

Chajnaetoitus, 272

Chameleo, 196

Chameleon, 196

Champse, 176

ehamses, Crocodilus, 177

changannvcnsis, Boulengerula, 306

chanleri, Mabuya, 211

chanleri, Simoeephalus, 254

chaperi, Riopa, 215

chapini, Raiia, 339

c-hapiui, Rana f., 339, 340

Chaunus, 309

chaus. Gecko, 183

Chelona, 164

Chcloue, 165

Chelonia, 163, 164, 165

CHELONIIDAE, 164

Chelyra, 164

clienonii, Dendrophis, 261

Ghersea, 300

Chersine, 169

Chersinella, 169

Chersophis, 300

Chersus, 169

chcvalieri, Rouleophis, 285

Chiamela, 213

Chilolepis, 257

Chilophryne, 309

Chilorhinophis, 284

Chioninia, 208

Chiriudia, 237

Chiromantis, 314

ehitialensis, Phrynobatiachus, 350
Chloroechis, 159, 303
cliloroechis, Vipeia, 159, 303, 304
Cklorophis, 259, 260
Chloroscopus, 232
chlorostigma, Varanus, 235
Chorisodon, 278
Choristocalamus, 281
Choristodon, 281
ehristyi, Aparallactus, 286
e-hristyi, Boulengeriiia, 291
rhristyi, Hylanibates, 318
ehristyi, Leptopelis n., 318
ehristyi, Miodon, 283
ehristyi, Miodon c, 283
ehristyi, Rana, 340
Chrysolamprus, 229
chrysoprasinus, Trypheropsis, 337
Cieigna, 225

cinctum, CoiJeglossiiin, 20S
Cinixys, 170
Cinothorax, 170
eiternii, Coluber, 258
citernii, Hemidactylus, 184
eitrinus, Hyperolius, 332
eitrinus, Hyperolius c, 332
Cladophis, 274
clathrotis, Lygosoma, 216
cliff ordii. Coluber, 257
Clinotarsus, 337
elivii, Xenopus, 307
aothelaps, 296
Clotho, 301
clotho, Cobra, 301
Clothonia, 248
Cnemaspis, 180
coarti, Atractaspis, 298
Cobra, 159, 301
eoecotis, Hyperolius, 332
toctaei, Hemidactylus, 181
collare, Miodon, 283
collaris, Cereocalamus, 285, 2>S7
collaris, Miodon e., 283

LOVERIDGK: E. AFRICAN' REPTILES AND AMPHIBIAXS

IX

Colobopus, 191
colonorum, Agama, 192, 194
Colpochelys, 168
Coluber, 2.")7, 270
COLUBRIDAE, 249, 268
colubrina, Dendrophis, 273
COLUBRIXAE, 249
eoniorensis, Euprepes, 210
comorensis, Mabuya in., 209, 210
r-omorensis, Tvphlops, 244
nompactilis, Bufo, 309, 310
complanatus, Crocodilus, 177
compressicauda, Lygosoma, 217
coucolor, Aparallactus 1., 286
concolor, Choristodon, 281
concolor, Hyperolius, 332
I'oncolor, Leptopelis, 318
concolor, Uriechis, 280
condanarus, Coluber, 278, 279
congestus, Onycliocephalus, 242
congica, Agama c, 194
congica, Atractaspis, 296
congicus, Aparallactus, 286
congicus, MegaJixalus f., 322
congicus, Scincodipus, 220
congicus, Tvphlops, 242
conica, Boa, 248

conjuncta, Leptotyphlops e., 246
conjunctum, Stenostoma, 246
conuali, Rhamphiophis, 277
conradsi, Atractaspis, 297
conradti, Lygodactylus, 187
Constrictor, 248
I'onstrictor, Coluber, 257
controversa, Thalassochelys, 168
Copeglossum, 208
Coracodichus, 350, 351
CORDYLIDAE, 223
Cordylus, 223
cordylus, Laeerta, 223
coriacea, Dermoehelys, 164
coriacea, Testudo, 164
Coriudo, 164
coronata, Calainaria, 258, 259

coronatus, Meizodon, 259
coronis, Mesocoronis, 300
corpulentum, Brachycrauion, 296
corticata, Thalassochelys, 167
Coryphodon, 257
cotti, Rana, 338, 341
cowani, Microscalabotes, 1S6
cowieii. Alligator, 177
Cranopsis, 310
Crassonota, 197
crawshayi, Naja n., 292
crepidianus, Phelsuma, 190
Orepidius, 310
Cricochalcis, 226
cristatus, Chamaeico, 197
croeea, Laeerta, 229
Crocodilus, 176
crocodilus, Laeerta, 176, 177
CROCODYLIA, 176
CROCODYLIDAE, 176
Crocodylus, 176
Crotaphitis, 337
Crotaphopeltis, 271
cruciger, Gekko, 185
cruentatus, Bufo, 308
Crypsidonius, 264
Cryptoblephaiiun, 218
Cryptoblepharus, 218
CRYPTODIRA, 163
Cryptopodus, 172
Cryptopus, 172
Cryptothylax, 316
cryptotis, Rana d., 345
cubitoalbus, Hylanibates, 318
cucullatus, Chamaeleo, 197
cuneiformis, Cadmus, 266
currori, Feylinia, 222
Cursoria, 249
curtus. Python, 248
Cusoria, 249
cutipora, Rana, 336
cyanogaster, Agama, 195
eyanogaster, Stellio, 193, 195
cyanophlyctis, Rana, 33<)

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Cyclanorbis, 172
Qv-clodei-ma, 172
CVclosaurus, 192
Cylindrurus, 235
Cyneosaura, 197
C.>Tiisca, 237

cynnamomeus, Hylambates, 317
cynodou, Dipsaa, 268
Cynodontophis, 283
Gyoclosaurus, 192
dabagae, Chamaeleon w., 205
Daboia, 300
Dactylethra, 307
damaranus, Euprepes, 212
damarensis, PelomtMiusa g., 174
darlingi, Honiopus, 170

DASYPELTINAE, 2S7
Dasypcltis, 162, 287, 288
(leeipiens, Hyperolius, 330
decosteri, Elapsoidea, 290
deeosteri, Elapsoidea s., 290
dofilippii, Caiisus, 299
defilippii, Heterodon, 299
degeni, CrotaphopeltLs, 272
degeni, Leptodira, 272
degrijisi, Gonionotophis, 255
Deirodon, 288
delalandii, Euprepes, 208
delalandii, Lacerta, 230
delalandii, Pyxicephalus, 337, 344
delalandii, Eana d., 344, 345
Dendraspis, 293
Dendroaspis, 293
Dendrobates, 308
dendrobates, Arthroleptis, 348
dendrobates, Phrj-nobatrachus, 348
Dendroeehis, 293
dendrophila, Dipsas, 268
Dendrovaranus, 235
denhardtii, Boulengerula, 305
dentatus, Sternotherus, 175
depressa, Chelonia, 166
depressus, Hemidactylus, 181
derbianus, Sternotherus, 175

deremensis, Chamaeleo, 205
deremeusis, Cliamaeleon, 205
Dermatochelys, 164
DERMOCHELYIDAE, 163
Dermochelys, 163
deserticola, Kassina, 321
devilliersi, Pelomedusa g., 174
dewittei, Siaphos, 216, 217
dhara, Coluber, 270
dhara, Telescopus d., 270
Diaphorotyphlops, 240
Diceros, 196

dickersoni, Gonatodes, 180
Dicranosaura, 197
Dicroglossus, 337
diesneri, Mabuia, 211
Dilepis, 197

dilepis, Chamaeleo, 197, 200
dilepis, Chamaleo d., 199
dimidiatus, Rhinocalamus, 282
Dinodipsas, 298
Dinophis, 293
Dioptroblepharis, 232
Diphalus, 237
DIPLASIOCOELA, 314
Diplodactylus, 189
Dipsadoboa, 269, 271
DIPSADOMORPHINAE, 268
Dipsadomorphus, 268
dipsas, Herpetodryas, 257
Dipsina, 276
Dipsoglyphophis, 271
Dirodon, 288
Discodaetylus, 189
discodactylus, Hyperolius, 326
discodaetylus, Phrynobatraehus, 347
DISCOGLOSSIDAE, 307
disjuncta, Pelomedusa g., 173, 174
disparilis, Diaphorotyphlops, 240
Dispholidus, 273
distanti, Agama h., 193
distanti, Leptotyphlops, 246
distanti, Leptotyphlops e., 246
Distichurus, 298

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

XI

Docidophryne, 309
dodomae, Agama a., 195
dodomae, Agama 1., 195
Dolichophis, 257
dolloi, Aparallactua, 286
dominiceusis, Mabuya, 208
dorbignyi, Bufo, 309
doriae, Ceramodactylus, 179
doriai, Latastia, 237
dorsalis, Afrixalus f., 322
dorsalis, Eremias, 232
dorsalis, Hyperolius, 322
dorsalis, Leptophis, 262
dorsalis, Philotliamnus s., 262
dorsalis, A'ipera, 298
dorsimaculatus, MegalLxalus, 322
Doryura, 181
Dracaena, 234
dracaena, Lacerta, 235
drapiezii, Dipsas, 268
Dromophis, 159, 277, 278
Dromoplectus, 310
Dryomedusa, 273
Duberria, 266
duberria. Coluber, 266
dubia, Clielonia, 168
dubia, Phelsuma d., 190
dubius, Pachydaetylus, 190
dubius, Typhlops, 240, 242
dugesii, Lacerta, 229
dulcis, E«na, 246
Dumerilia, 220
dumerili, Euprepes, 215
dumerilii, Coronella, 256
dumerilii. Monitor, 235
dussumieri, Chelonia, 168
dutoiti, Arthroleptides, 336
Ebeuavia, 190, 359
Echidna, 301
Echidnoides, 300
ochinata, Lacerta, 230
Eehis, 303
echis, Vipera, 303
edulis, Pyxicephalus, 337, 344

edulis, Rana, 344
cdwarsiana, Lacerta, 228
eggeli, Scelotes, 219, 220
Elabites, 208
Elapechis, 290
Elaphe, 257
Elaphrodytes, 295
Elaphromantis, 31C
Elaphropus, 229
ELAPIDAE, 290
Elapomorphus, 284
Elaposchema, 282
Elaps, 291
Elapsoidea, 290, 291
Elasmodaetylus, 191
elegans, Acontias, 222, 223
elegans, Arthroleptis, 353
elegans, Coluber, 278
elegans, Coronella, 259
elegans, Cusoria, 249
elegans, Euprepes, 214
elegans, Feylinia c, 223
elegans, Miculia, 218
elegans, Pachydaetylus, 191
elegans, Stenodactylus, 179
elegans, Testiido, 169
elegans, Tupinambis, 236
elegans, Vipera, 300
elegantissimus, Coluber, 266
elegantissimus, MLke, 279
elgonensis, Cnemaspis a., 180
elgonensis, Rhamnophis a., 264
elgonensis, Thrasops a., 264
elgonis, Agama, 195
elgonis, Agama a., 195
ellenbergeri, Chamaetortus a., 272
ellenbergeri, Mabuya s., 213, 272
ellioti, Chamaeleo b., 200
ellioti, Chamaeleon, 200
elongata, Caouana, 167
elongata, Dasypeltis, 289
elongata, Eremonia, 167
embuensis, Chamaeleon a., 203
emini, Ahaetulla, 261

xn

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

enimi, Glaueonia, 247

emini, Leptotyphlops e., 247

eniini, Nucras, 230

Eminophis, 159

Empagusia, 235

empusa, Peltaphryne, 309

eiiantiodaetylus, Hylambates, 315

Endodroniius, 228

Engystoma, 357

ENGYSTOMATIDAE, 355

Ensirostris, 197

Epictia, 246

Epidalea, 309

cpioticus, Crepidius, 310

Eremias, 232

Eremioplanis, 192

Ereniioscopus, 232

Eremonia, 167

Eretmoclielys, 166

Erizia, 197

erlangeri, Hemidactylus, 185

erlaiigcri, Eana, 341

orosa, Kinixys, 171

crosa, Testudo, 170, 171

erythraea, Hyla, 336

Eryx, 162, 248

esehiichtii, Typhlops, 239, 241

esculenta, Caretta, 165

esculenta, Rana, 336, 337

etiennei, Chamaeleo g., 199

EUBLEPHARIDAE, 178

Eucephalus, 245

Euchelonia, 165

Euehelys, 165

Eudioptra, 232

Eudipsas, 268

Eugnathus, 250

Eugong3''lus, 214

Euleptes, 189

Euniecia, 213, 214

Euphlyctis, 336

Euprepiosaurus, 235

Euprepis, 208

I'uprootus, Typhlops, 244

Eurhina, 309
Eurhous, 182

europaeus, Phyllodactyhis, 189
Eurystephus, 296
Eutropis, 208
Evoluticauda, 206
ewerbeeki, Amphisbaeua, 238
ewerbecki, Cliirindia, 238
exanthematiea, Lacerta, 23.5
t'xanthematicus, Varanus e., 236
oxcentricus, Typhlops, 241
oxceutrieus, Typhlops s., 241
oxcubitor, Chamaeleo f., 203
t'xcubitor, Chamaeleon t., 203
I'allax, Atractaspis, 296, 297
fallax, Taibophis, 270
fasciata, Dasypeltis, 288, 28!i
fasciata, Dipsas, 268
fasciata, Grayia, 268
fasciata, Hoimonota, 182
fasciata, Pelamis p., 296
fasciata, Platypholis, 191
fasciata, Platypholis p., 191
fasciatus, Chiromantis, 316
fasciatus, Dasypeltis, 289
fasciatus, Hemidactylus f., 181
fasciatus, Strongylopus, 336
fasciolata, Dasypeltis s., 289
fasciolatus, Coluber, 258
fastidiosus, Cranopsis, 310
fenestratus, Diphalus, 237
feniandi, Riopa, 213, 214
fernandi, Tiliqua, 214
fevniquei, Eappia, 329
ferrandii, Lygosoma, 21;")
Feylinia, 222
fiechteri, Glaueonia, 247
fiechteri, Leptotyphlops, 247, 3.")0
fiechteri, Rana, 338
FIRMISTERXIA, 314
fischeri, Chamaeleo f., 204, 359
fischeri, Chamaeleon, 204
fischeri, Dipsas, 269
fischeri, Zamenis, 259

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS xiii

fitzingeri, Notopholis, 228
fitzsimonsi, Gerrhosaurus f ., 225
flaveseens, Monitor, 235
flavigula, Pyxicephalus, 344
flavigularis, Dendrophis, 263
flavigularis, Gerrhosaurus, 224
flavitorques, Aparallactus, 285
flavoguttatus, Hyperolius, 328, 335
flavomaculatus, Hyperolius, 318
flavomaculatus, Leptopelis, 318
flavoviridis, Hyperolius, 332
fletcheri, Typhlops, 245
florulentulus, Coluber, 258
florulentulus, Coluber f ., 258
floweri, Eana, 338, 341
Fordia, 171
for mesa, Ohelonia, 166
fornasini, Afrixalus f., 322, 325
fornasini, Euchnemis, 322
fragilis, Heniidactylus, 184
francei, Arthroleptis a., 352
franklini, Oriychocephalus, 240
frenatus, Cycloderma, 172
frenatus, Hemidactylus, 181, 183
frenatus, Xonurus, 223
f riedericianus, Adolfus, 228
frontatus, Crocodilus, 178
fiilleborni, Chamaeleo, 205
fiilleborni, Chamaeleon, 205
fuelleborni, Hyperolius, 328
fiilleborni, Eana, 343
fiilleborni, Eana f., 343
fuliginosa, Amphisbaena, 237
fuliginosus, Boaedon f ., 251
fuliginosus, Lycodon, 250, 251
fulvicollis, Microsonia, 283
fulvicollis, IVIiodon, 283
fulvovittatus, Afrixalus f ., 323
funereus, Bufo, 311
Furcifer, 196
fuscigula, Eana, 340
fuscigula f., 389, 340
fuscorosea, Coronella s., 258
fuscum, Trachischum, 159

fuscum, Triglyphodon, 268
fuscus, Dipsas, 268
gabonensis, Elapomorphus, 283
gabonensis, Miodon, 283
gabonica, Bitis g., 302
gabonica. Echidna, 302
galamensis, Eana, 339
galamensis, Eana g., 338
galeata, Testudo, 173
galloti, Lacerta, 229, 230
Gallotia, 229, 230
gambiense, Lyeophidion c, 253
gardineri, Hemidactylus, 186
garmani, Bufo, 310
garnotii, Hemidactylus, 181
gasconi, Pelomedusa, 173
Gastropholis, 226
Gastropyxis, 263
gecko. Gecko, 181
gehafie, Pentonyx, 173
geitje, Lacerta, 191
GEKKONIDAE, 178, 179
gemmatus. Coluber, 279
gemonensis, Natrix, 257
Geocalamus, 238
Geochelone, 168, 169
Geodipsas, 255, 268
geometrica, Testudo, 169
geometricus, Lycodon, 250
geiardi, Apostolepis, 284
Gerrhopilus, 239
Gerrhopygus, 189
gerrhopygus, Diplodactylus, 189
GEEEHOSAUEIDAE, 223
Gerrhosaurus, 224
giardi, Grayia, 256
gibbosa, Eana, 356, 357
gibbosus, Breviceps, 356
gierrai, Typhlops, 242
gierrai, Typhlops p., 242
Gigantorana, 338
gigas, Caretta, 168
glaber, Hetevolepis, 253
glaber, Tropidonotus, 267

x\y

BULLETIN : MUSEUM OP^ COMPARATIVE ZOOLOGY

^landicolor, Hyperolius, 334, 359
Glaniolestes, 267
Glauconia, 246
Globiceps, Dipsas, 269
slobosa, Sehoutedenella, 351
glohulosus, Chaimus, 309
(ilypliolycus, 250
ftootzei, Chainaeleo g., 202
goetzei, Cliamaeleon, 202
goetzei, Hj'perolius, 325
ffoldii, Naia, 293
goldii, Pseudohaje, 293
Roliath, Raiia, 338
gollmeri, Rana, 337
Goiigvloniorphus, 219, 220
(jongylophis, 248
gouldii, Hydrosaurus, 235
gracilis, Aspidorhinus, 232
gracilis, Chamaeleo, 198
gracilis, Chamaeleo g., 198
gracilis, Chlorophis, 262
gracilis, Leptophis, 263
gracilis, Phrynobatrachus n., 348
gracilis, Typhlops, 243, 245
gracillima, Ahaetulla, 261
graeca, Testudo, 16<S, 169
gramineus, Leptopelis, 316, 359
gramineus, Megalixalus, 316
grandiceps, Leptopelis, 320
grandis, Gerrhosaurus, 224
grandis, Gerrhosaurus m., 224
gvaiiosum, Engystoma, 357
graiitii, Euprepes, 213
gianulata, Rappia, 333
granulosus, Bufo, 309
granulosus, Chamaeleo, 198
granulosus, Hyperolius, 332
giaueri, Aparallactus, 286
graueri, Arthroleptis, 348
graueri, Chamaeleon, 205
gi aueri, Chamaeleon b., 200
graueri, Hyperolius, 330
graueri, Leptopelis, 319
graueri, Lygo.soma, 217

graueri, Lygosoma g., 217
graueri, Mabuia m., 209
graueri, Miodon, 284
graueri, Miodon c, 284
graueri, Phrynobatrachus, 348
graueri, Typhlops, 243
grayanus, Blepliarostercs, 218
Grayia, 267
grayii, Trioceros, 196
gregorii, Agama, 196
gregorii, Dermophis. 30")
gregorii, Schistometopum, 305
greshoffi, Hylainbates, 316
greyii, Menetia, 218
gribiuguiensis, Rana, 340
gri])inguicnsis, Rana o., 340
griseus, Lygodactylus p., 188
griseus, Tupinambis, 234
Grobbenia, 254
gromieri, Lygosoma, 217
grotei, Lygodactylus, 187
giotei, Lygodactylus g., 187
groutii, Stenostoma. 246
griitzneri, Euprepes, 213
Grypotyphlops, 240
gueinzii, Heterolepis, 254
giintheri, Amplorhinus, 275
guentheri, Aparallactus, 287
giintherii, Elapsoidea, 290
guntherii, Elapsoidea s., 290
guentheri. Holaspis, 227
guentheri, Hypogeophis, 305
giintheri, Philothanmus, 261
guentheri, Tarbophis, 270
guineensis, Bufo, 310
guiueeusis, Chiromantis, 315
guineensis, Crotaphopeltis d., 271
guineensis, Eremias, 233
guineensis, Hemidactylus, 184
guineensis, Hemisus, 355
guineensis, Hemisus m., 355
guineensis, Leptodira, 271
guineensis, Riopa s., 215
guttatolineatus, Hyperolius m., 334

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

XV

guttatum, Engystoma, 354
guttatus, Bufo, 309
guttatus, Lycodon, 250
guttatus, Stenodactylus, 179
gutterosus, Phrynobatiachus, 349
guttolineatus, Hyperolius, 334, 359
guttulata, Lacerta, 232
gutturalis, Bucephalus, 273
gutturalis, Hemidactylus, 188
gutturalis, Lygodactylus p., 188
Guyomarchia, 285
GYMNOPHIONA, 305
Gymnopus, 171
liaasi, Leptopelis b., 317
liackarsi, Lygosoma m., 217
liaemachatea, Vipera, 291
haematiticus, Bufo, 309
Haemorrhois, 257
Hagria, 213

liailensis, Phrynobatrachus, 340
haje, Coluber, 275, 291
haje, Naja, 275
haje, Naja h., 291
Halcrosia, 178
Halichelys, 167
hammondii, Dinophis, 293
Hapsidophrys, 263
harlani, Plestiodon, 214
Heleionomus, 248
Heliobolus, 232
Heliophilus, 230
helleri, Siaphos m., 217
Hemidactylus, 181
Hemimantis, 350
Hemirhagerrhis, 275
Hemisus, 214, 335, 354
Hemorrhois, 257
hemptinnei, Scelotes t., 221
hemptinnei, Sepsina, 221
Heptathyra, 172
Herinia, 208, 219, 220
Herpetaethiops, 259
Herpetosaura, 219, 220
Herpetoseps, 220

Herpysmostes, 166

HeterLxalua, 321

heterodermus, Cliloropliis, 259, 260

heterolepidota, Ahaetulla, 261

heterolepidotus, Philothamnus, 261,

262
Heterolepis, 254
Heteronotus, 267
Heteropelis, 316
Heterophis, 298
heterurus, Ophis, 272
hexacera, Vipera, 303
hexadactyla, Rana, 336
hexaphractos, Crocodilus, 177
hexaspis, Hemidactylus, 183
hieroglyphiea. Boa, 248
hieroglyphica, Rana, 345
Hierophis, 257
hildebrandti, Mabuia q., 209
Hildebrandtia, 337
hildebrandtii, Ablabes, 275
liildebrandtii, Atractaspis, 281
liildebrandtii, Euprepes, 212
liindii, Vipera, 300
hippocrepis. Coluber, 257
hippocrepis, Dipsas, 272
hispanicus, Psaminosaurus, 228
hispida, Agama, 193
hispida, Atheris, 304
hispida, Lacerta, 192
lioehneli, Eremias, 233
hohnelii, Chamaeleo b., 201
hohnelii, Chamaeleon, 201
Holaspis, 227
Holodactylus, 178
Holuropholis, 250
Homalosoma, 266
homeyeri, Ablabes, 280
Homodactylus, 191, 225
Homopholis, 190
Hoplobatrachus, 337
lioplogaster, Ahaetulla, 260
hoplogaster, Chlorophis, 260
hoplogaster, Philothamnus, 26U

XVI

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

HoplopliTjTie, 358
Hoplopodion, 181
Hoplopodium, 182
Hoimonotus, 253
horsfieldi, Testudo, 169
horstokii, Lycodon, 251, 252
Hortulia, 248

hotamboeia, Coronella, 271
liotamboeia, Crotaphopeltis h., 271,

272, 273
hottentotus, Typhlops, 241
houttuyni, Python, 248
houyi, Eiopa, 215
liugyi, Vipera, 300
humbo, TjTphlops, 241
hydraletis, Eana, 344
HYDROPHIIDAE, 295
Hydrophylax, 336
Hydrosaurus, 235
Hydrus, 295
Hylambates, 320
Hylaplesia, 308
Hylarana, 336
Hylarthroleptis, 346
Hylorana, 336
Hypapistes, 247

HYPEROLIIDAE, 163, 314, 335
Hyperolius, 161, 314, 324
hypsirhinoides, Nerophidion, 250
iberus, TrigonopMs, 270
Ibiba, 159, 268
Ichnotropis, 159, 233, 234
idae, Rana, 342
imbricata, Eretmochelys, 166
imbricata, Testudo, 166
incanescens, Gecko, 185
inconspicuous, Typhlops, 244
incornutus, Chamaeleo, 202
incornutus, Chamaeleon, 202
indicus, Tupinambis, 235
Indovaranus, 235
inexpectatus, Phyllodactylus, 189
inflata, Vipera, 302
iiifralineata, Mabuia c, 210

inf ralineatus, Taehymenis, 255
inguinalis, Arthroleptis, 352
inliambanensis, Dendrobates, 355
inornata, Coronella, 259
inornatus, Dipsas, 272
inornatus, Hemidactylus, 183
inornatus, Lithophilus, 220
intermedia, Cassina s., 321
intermedia, Gerrhosaurus f., 225
intermedia, Typhlops 1., 242
intermedius, Crocodilus, 176
intermedius, Dendraspis, 294
intermedius, Psammophis s., 279
intermixta, Naja, 291
intumescens, Coluber, 302
inunguis, Anoplopus, 191
inuuguis, Ebenavia, 190
inunguis. Gecko, 191
ionidesi, Amblyodipsas k., 282
ionidesi, Anclocranium, 237
ionidesi, Brookesia, 207
ionidesi, Brookesia b., 207
ionidesii, Amphisbaena, 238
ipianae, Hyperolius, 323
irregularis, Atractaspis, 296
irregularis, Atractaspis i., 297
irregularis, Coluber, 159, 261. 268
irregularis, Elaps, 296
irregularis, Hyperolius, 327
irregularis, Mabuia, 208, 212
irregularis, Mabuya, 212
iri'egularis, Philothamnus i., 261, 262
irregularis, Psammophis, 279
irrorata, Arthroleptis n., 348
irroratus. Coluber, 252
isabellina, Phayrea, 278
isabellinus, Chamaeleo d., 199
Isodactylus, 193
isodactylus, Eumeees, 214
isolepis, Hemidactylus, 182
isselii, Euprepes, 212
jacksoni, Aparallaetus j., 28(i
jaeksoni, Lacerta, 229, 230
jacksoni, Lyeophidium, 252

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS xvii

jacksonii, Causus, 299
Jackson ii, Chamaeleo, 205
jacksonii, Chamaeleon, 205
jacksonii, Thrasops, 264
jacksonii, Thrasops j., 264, 273
jacksonii, Uriechis, 286
jamesoni, Elaps, 293
janenschi, Hylarthroleptis, 346
japonica, Testudo, 165
jardinii, Bucephalus, 273
javanicus, Hemidactylus, 181, 183
jeanneli, Mabuia, 208, 212
johnii, Erj-x, 248
johnsoni, CrocodUus, 176
johnstoni, Chamaeleo j., 205
johnstoni, Chamaeleon, 205
johnstoni, Crocodilus, 176
johnstoni, Hylambates, 318
johnstoni, Latastia, 231
jonesi, Cordylus c, 223
jordani, KinLxys, 171
jouberti, Pelomedusa, 175
jubalis. Python, 248
kachowskii, Chiromantis, 315
kageleri, Crotaphopeltis h., 271
kaimosae, Agama a., 195
kaimosae, Dendraspis j., 294
kaimosae, Demlroaspis j., 294
kaimosae, Typhlops, 243
Kakophrynus, 354
kandti. Hyper olius, 327
karissimbensis, Leptopelis, 319
karissimbiensis, Hyperolius, 327
Kassina, 320
Kassimila, 321
katangae, Atractaspis, 298
kaznakovi, Vipera, 300
kelaarti, Bufo, 309
kelleri, Chiromantis, 315
kelleri, Chiromantis p., 315, 316
kelleri, Hemirhagerrhis, 275
kempi, Colpoehelys, 168
keniensis. Coluber, 258
keniensis, Lygodactylus p., 188

keniensis, Mabuya b., 211, 212
keniensis, Phrynobatrachus, 349
kerstenii, Brookesia k., 206
kerstenii, Chamaeleo, 207
kibonotensis, Lacerta j., 230
kidweJli, Latastia, 231
kilimense, Lygosoma, 216
kilimensis, Lygosoma, 216
kilosae, Nucras b., 231
kUosae, Lucras b., 231
kinangopensis, Phrynobatrachus, 347
kinetensis, Chamaeleo b., 200
Kinrxys, 170
Kinothorax, 170
kirkii, Ahaetulla, 262
kirkii, Chamaeleon p., 200
kirkii, ScoIe<;omorphus, 306, 307
kirkii, Scolecomorphus k., 307
kirtlaudii, Leptophis, 274
kirtlaudii, Thelotornis k., 274
kisoloensis, Bufo r., 311
kivuensis, Dispholidus t., 274
kivuensis, Hyperolius, 326, 332
kivuensis, Leptopelis, 319
kleebergi, Typlilops, 245
koehli, Hyperolius, 326, 327
komodoensis, Varanus, 235
kraussi, Onychochelys, 166
kraussi, Typhlops, 242
kreffti, Callulina, 356
krefftii, Phrynobatrachus, 347
krevirostris, Crocodilus, 17G
kuhlii, Kana, 336
kutuense, Lygosoma, 217, 359
kutuensis, Lygosoma, 217
kuvangeusis, Cassiniopsis, 321
kwidjwiensis, Hyperolius, 326
kwidjwiensis, Mabuia m., 209
labiatus, Trionyx, 172
Lacerta, 229
LACERTIDAE, 226
LACERTILIA, 178
lachesis, Cobra, 159, 301
lachrymata, Chelonia, 165

XVlll

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

lacunosus, Crocodilus, 177
laeviceps, Athens, 304
laevigatus, Chamaeleo, 198
laevigatus, Euprepes, 212
laevis, Bufo, 307
laevis, Holaspis g., 227
laevissinia, Natrix, 249
lalandii, Dispholidus, 273
lalandii, Lacerta, 230
lalandii, Typhlops, 239, 240
lamani, Mehelya, 254
lamarrei, Campsodactylus, 213
Lampreremias, 232
lamuensis, Dasypeltis m., 289
lata, Chelonia, 166
Latastea, 300
latastei, Vipera, 300
Latasteopara, 300
Latastia, 231
laterale, Lissophis, 252
lateralis, Chamaeleo, 196
lateralis, Schismaderma, 309, 310
laterispinis, Chamaeleo, 202
laterispinis, Chamaeleon, 202
lateristriga, Clotho, 302
laticeps, Pappophis, 268
latifions, Bufo, 162, 311
latifrons, Hyperolius, 327
latirostris, Glauconia, 246
latirostris, Typhlops, 241
Laudaiia, 193
laurae, Lygodactylus, 187
ieachi, Anilios, 240
Leaehianus, Sternothaerus, 175
lebetimis. Coluber, 300
lecomtei, Oxybelis, 274
leikipiensis, Chamaeleon, 201
leioccphalus, Chamaeleon s., 198
Leionotus, 267
Leiurus, 181, 182
leleupi, Cacostemum, 354
ieuzi, Typhlops, 245
leonaidi, Hylambates, 320
looninus, Euprepes, 214

leopardinus, Psammophis s., 279
leopoldi, Melanocalamus, 283
Lepidochelys, 168
Lepidon, 291
Lepidothyris, 214
Leptoparius, 346
Leptopelis, 316
Leptophryne, 308
leptorhynchus, Crocodilus, 177
Leptosiaphos, 216
leptosomus, Afrixalus f., 322
leptosomus, Hyperolius, 323
LEPTOTYPHLOPIDAE, 239, 245
Leptotyphlops, 239, 245
leptus, Pelusios, 175
lepus, Chiromantis, 338
Lerista, 218

leschenaultii, Bufo, 309
leschenaultii, Hemidactylus, 182
leschenaultii, Eana, 336
lestradei, Typhlops, 240
lestradei, Typhlops b., 240
Letheobia, 240

leucopunctata, Hylambates, 317
leucosticta, Naja h., 293
leucostriata, Pelamis p., 296
leucura, Amphisbaena, 237
Levirana, 337
Liasis, 247

liberiensis, Onychocephalus, 242
liberiensis, Ophioproctes, 237
liberiensis, Python, 248
liehtensteinii, Aspidelaps, 298, 299
lichtensteinii, Causus, 299
liehtensteinii, Mesalina, 232
Ligonirostra, 265
limbricki, TjT)hlops, 244
Limnodytes, 330
Limnonaja, 290, 291
Limnonectes, 336
Limnophilus, 336
lindneri, Bufo, 312, 359
lineata, Chiamela, 213
lineata, Hapsidophrys, 263

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

XIX

linea.ta, Lerista, 218
lineatum, Boaedon, 250, 251
liiieatus, Acontias, 239
lineatus, Boaedon 1., 160
lineatus, Bothrophthalmus 1., 249
lineatus, Causus r., 298
lineatus, Dromophis, 278
lineatus, Dryophylax, 278
lineatus, Elaphis, 249
lineatus, Hapsidophrys, 263
lineatus, Typhlina, 240
lineatus, Typlilops, 239, 240
lineatus, Uriechis, 285
lineolata, Eminophis, 159
lineolata, Pseuderemias, 232
lineolatus. Coluber, 278
lineolatus, Dasypeltis, 288
lineolatus, Typhlops, 242
lineo-ocellata, Eremias, 232
lionotus, Agania, 194
lionotus, Agama a., 194
Liophallus, 268
Liosoma, 213, 214
Lissophis, 251
Lithophilus, 220
Li urns, 182

livingstonii, Aspidochelys, 172
liwalensis, Chilorhinophis c, 284
lobatsiana, Cinixys, 170
lonnbergi, Arthroleptis, 352, 353
lonnbergi, Bufo, 312
lonnbergi, Bufo 1., 312
longicauda, Leptotyphlops, 247
longicauda, Melanoseps a., 221
longicaudata, Lacerta, 231
longicaudata, Latastia 1., 231
longicaudum, Stenostoma, 247
longiceps, Hemidactylus, 183
longifrenatus, PMlothamnus i., 261
longissimum, Ophthalmidium, 240
longissimus, Elaphe, 257
Lophopholis, 182
Lophosaura, 197
LORICATA, 176

Loveridgea, 238
loveridgei, Atractaspis i., 297
loveridgei, Dasypeltis s., 288
loveridgei, Elapsoidea s., 290
loveridgei, Eryx c, 249
loveridgei, Eryx t., 249
loveridgei, Hyperolius s., 332
loveridgei, Meizodon, 258
loveridgei, Probreviceps m., 357
loveridgei, Ptyehadena, 343
loveridgei, Rana, 343
loveridgii, Megalixalus, 322
loveridgii, Testudo, 170
Loxodon, 257
lueani, Atheris, 304
lugubris, Lacerta, 232
lumbricalis, Anguis, 239, 240
lumbriciformis, Onychocephalus, 245
lunibricif ormis, Typhlops, 245
luuulatus, Aparallactus 1., 286
lunulatus, Uriechis, 285, 286
lutaria, Chelonia, 164
iuteus, Ophinectes, 295
lutrLx, Coluber, 266
lutzei, Gastropholis, 226
Lycodonomorphus, 249
Lycodontomorphus, 249
Lyeophidion, 251, 252
Lycophidium, 252
Lyeophydiou, 251
Lygodactylus, 186
Lygosoma, 213, 216, 220
Lygosoma gromieri, 217
lyra, Testudo, 164
niababiensis, Kinixys a., 171
mababiensis, Phrynobatraehus, 350
niababiensis, Phrynobatraehus u.,

350
mabouia. Gecko, 181, 185
mabouia, Hemidactylus, 160, 185
mabouya, Lacertus, 208
mabuia. Gecko, 181
niabuiiformis, Eiopa, 213, 214
Mabuiopsis, 208

XX

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Mabuya, 220

maerocephala, Ptychadena, 342
Maerocephalus, 268
macrodaetylus, Arthroleptis, 352
nmcrodactylus, Breviceps, 356
macrodaetylus, Hyperolius, 326
macrodaetylus, Probreviceps m., 356
macrolepidota, leliiiotropis, 234
macrolepis, Mancus, 226
macrolepis, Stenostoma, 246
Macrophis, 267

macropholis, Hemidactylus, 185
macropholis, Hemidactylus t., 185
macrops, Chiromantis, 315
macrops, Dasypeltis, 289
macrops, Oligolepis, 260
macrops, Philotliamnus, 260
macropus, Euchelys, 165
maeropus, Testudo, 165
macrorhynchum, Stenostoma, 246
Macrosoma, 278
macrotympanum, Eana, 345
Macrovipera, 300
macularius, Euprepes, 208
maculata, Hydrophis b., 295
maculatus, Boaedon, 251
maculatus, Distiehurus, 298
maculatus, Hylambates, 317, 320, 321
maculilabris, Euprepis, 209
maculilabris, Mabuya m., 209
maculosa, Chelonia, 165
maculosus, Megalixalus, 321
madagaseariensis, Crocodilus, 177
madarensis, Lacerta, 229
magretti, Raua, 338
major, Gerrhosaurus, 224
major, Gerrhosaurus m., 224
major, Mabuia m., 209
malabariea, Rana, 336
Malacochersus, 169
Maltzania, 337
mamba, Dendraspis, 294
Mancus, 226
mandanus, Hemidactylus, 186

mandensis, Typhlops, 241
mandera, Brookesia, 206
manni, Lygodactylus, 189
marakwetensis, Bufo r., 311, 359
margaritata, Lacerta, 229
margaritifer, Bufo, 309
margaritifer, Mabuya q., 208
margaritiferus, Coryphodon, 259
margiuata, Testudo, 169
marginatus, Crocodilus, 177
marginatus, Hylambates, 317
marginatus, Hyperolius, 325
niariae, Hyperolius, 328, 331
marina, Rana, 309
marina, Testudo, 165
marmorata, Chelonia, 16G
marmoratum, Engystoma, 354
marnioratus, Chaunus, 309
marmoratus, Hemisus m., 354
marmoratus, Hyperolius, 324
marmoratus, Pyxicephalus, 345
marshalli, Rhampholeon, 197
martensi, Latastia d., 231
martiensseni, Arthroleptides, 335,

336
martiensseni, Leptopelis, 319
marunguensis, Chamaeleo a., 198
mascarenieusis, Rana, 336, 337, 342
mascareniensis, Rana m., 341
massaiensis, Ablepharus, 219
massaiensis, Eumeces, 211
matengoensis, Melanoseps a., 222
matschiei, Chamaeleon, 204
mcphersoni, Melanelaps, 296
Mecistops, 176
Medaestia, 169
medici, Dasypeltis, 288
medici, Dasypeltis m., 289
medici, Dipsas, 289
medilineatus, Hylarthroleptis, 346
Meditoria, 240
Megablabes, 257
Megacliersine, 168, 169
Megalixalus, 321

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS

XXI

megalura, Euprepes, 211
iiiegalura, Mabuya, 211
megalurus, Ablepharus, 219
Alegemys, 165
Mehelya, 254
Meizodon, 258

inelancholicus, Anaxyrus, 309
Melanelaps, 296
MelanocaJamus, 283
melanocephala, Mierosaura, 197
melanocephalus, Cathetorhimis, 240
melanocrotaphus, Oxyropus, 272
melanoleuca, Naia h., 293
melanoleuca, Naja, 293
melanoleuca, Naja n., 292
melanophthalmus, Hyperolius, 328
Melanoseps, 219, 221
melanostigma, Dendrophis, 262
melanozostus, Bothrophthalmus, 249
melanura, Laudakia, 193
iiieleagris, Anguis, 222
meleagris, Calamaris, 264
meleagris, Lycophidion, 252
meleagris, Lycophidium, 252
meleagris, Lygosoma, 216, 217
meleagris, Rhiaocalamus, 282
mellaudii, Calamelaps, 281
melleri, Chamaeleo, 202
melleri, Ensirostris, 197, 202
menestriesii, Ablepharus, 218
Menetia, 218
mentalis, Boaedon, 251
mercatorius, Hemidactylus, 186
mereuralis, Sphargis, 164
meridionalis, Tarbophis s., 270
merkeri, Glauconia, 246
merremii, Euprepes, 208
mertensi, Chamaeleo a., 198
merumontana, Eana, 343
merumontana, Rana f ., 343
Mesalina, 232
Mesocoronis, 300
Mesohorinis, 300
Mesovipera, 300

methneri, Arthroleptis, 353
methneri, Spelaeophryne, 356
Metopophis, 285
Metoporhina, 252
micranotis, Bufo, 313
micranotis, Bufo m., 313
Micrarthroleptis, 351
Micrelaps, 282
MICEOHYLIDAE, 350
Microbatrachus, 346
Microbatrachella, 346
Microblepharis, 218
micioc-ephalus, Typhlops, 244
Microdactylus, 181
MICEOHYLIDAE, 314, 355, 356
niicrolepidota, Atractaspis, 297
mierolepidota, Atractaspis m., 296,

297
mierolepis, Eumec«s, 214
iiiicrolepis, Spelaerosophis, 257
microphthaluia, Calaniaria, 282
microps, Gonionotophis, 255
uiicrops, Hyperolius, 333
microps, Phrynomerus b., 356
Mierosaura, 197
Microsealabotes, 186
Microsoma, 283
microstictus, Monitor, 236
microstictus, Vai-anus, 236
microstictus, Varanus e., 236
Miculia, 218
iiiigiurtina, Eana o., 341
Migiurtinophis, 270
Mike, 279

milleti, Typhlops, 242
inilnei, Hyperolius, 333, 334
minor, Agama, 193
minutus, Arthroleptis, 349
minutus, Phrynobatrachus, 349
Miodon, 283

miolepis, Calamelaps, 282
miopropus, Chamaesaura, 226
niitchelli, Breviceps, 357
mlanjensis, Mabuya b., 212

XXll

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Mochlus, 213, 214
mocquardi, Buf o, 313
mocquardi, Riopa, 215
Moequardia, 320
modesta, Kassina, 321
modesta, Riopa m., 213, 215
inodestus, Aparallactus, 285
modestus, Bunocnemis, 182
inodestus, Elapops, 285
modestus, Geocalamus, 238, 239
modestus, Hemidactylus, 182
modestus, Hormonotus, 253
modestus, Lampropliis, 253
modestus, Sepacoutias, 214, 215
nioebiusi, Elapechis, 290
moliasicus, Hyperolius, 327
Molinia, 176
molurus. Coluber, 248
mombasieus, Lygodactylus p., 188
moniliger, Coluber, 279, 280
Monitor, 234

monitor, Lacerta, 234, 235
Mouodactylus, 226
Monodiastema, 278
montana, Agama m., 194
montana, Rappia, 326
montanus, Hyperolius, 326
monticola, Hyperolius, 327, 328
monticola, Phrynobatrachus, 345
moorii, Phrynobatrachus, 350
moreaui, Bedriagaia, 227
Morelia, 247

moreoticus, Algyroides, 228
Morethia, 218
mortoni, Trionyx, 172
inosaiea, Bufo, 310
mossambica, Agama, 193, 194
mossambica, Agama m., 193
mossambica, Coronella s., 259
mossambica, Lygodactylus c, 187
mossambica, Naja, 292
mossambica, Psammophis s., 279
mossambica, Rana m., 341
mossambicana, Thelotornis k., 274

mossambicensis, Pelomedusa, 174
mossambicus, Breviceps, 357
mossambicus, Dasypeltis s., 288
mossambicus, Thrasops j., 274
Motinia, 176

mozambiea, Pelomedusa, 174
mpwapwaensis, Amphisbaena, 238
mucronatus, Acanthodactylus, 232
mueruso, Onychocephalus, 241
mucruso, Typhlops s., 241
muelleri, Dactylethra, 308
miiUeri, Micrelaps, 282
muelleri, Phaneropis, 218
muelleri, Xenopus, 308
multicolor, Hyperolius, 327
multifasciatus, Hyperolius, 323
multifasciatus, Scincus, 208
multilineatus, Gerrhosaurus, 225
multilineatus, Typhlops, 239
multimaculata, Coronella, 276
multimaculata, Lycophidium c, 252
multimaculatus, Amplorhinus, 275
multiscutata, Chelonia, 168
multiscutatus, Crocodilus, 177
multisquamis, Psammophylax t., 276
multisquamis, Trimerorhinus t., 276
multituberculatus, Chamaeleo f ., 204
multituberculatus, Chamaeleon f.,

204
muralis, Seps, 229
mutabilis, Agama, 192, 193
mwanzae, Agama 1., 195
mwanzae, Agama p., 195
mwanzae, Rana, 344
Mydas, 164
mydas, Chelonia, 165
mydas, Testudo, 164, 165
Mydasea, 165

Naia, 291

nairobiensis, Bufo 1., 312, 313

Naja, 291

naja. Coluber, 291

namaquensis, Chamaeleo, 197

LOVERIDQE: E. APKICAN REPTILES AND AMPHIBIANS

XXlll

Nannophryne, 310
nanuni, Cacosternum, 353, 354
nanus, Paraithroleptis, 351
naricus, Oxyrliynclms, 309
nasalis, Causus, 299
nasicornis, Bitis, 302
nasicornis, Caretta, 165
nasicornis, Coluber, 302
nasicornis, Testudo, 165, 167
nasus, Brookesia, 206
nasuta, Caretta, 167
nasuta, Meditoria, 240
nasutus, Chamaeleo, 197
nasutus, Hyperolius, 333
iiatalensis, Ilylambates, 318
nat^leusis, Kinixys, 171
natalensis, Phrynobatrachus, 348,

349
natalensis, Python, 248
natalensis, Stenorhynchus, 346, 348
Natalobatrachus, 346
Natriciteres, 255, 256, 268
nchisiensis, Brookesia, 207
Xectophrynoides, 314
neglectus, Chlorophis, 260
noglectus, Philothamnus, 260
Nerophidion, 250
neumanni, Eremias, 233
neumanni, Philochortus, 227
neuricatenata, Pelamis p., 296
neurileucura, Pelamis p., 296
Xeusteropliis, 249
neuAviedi, Miodon, 283
neuwiedii, Mierosoma, 283
newtonii, Rana, 342
ngamiensis, Lygodactylus c, 187
ngoriensis, Hyperolius, 325
niangarae, Calamelaps, 281
niger, Crocodilus, 176
niger, Sternotherus, 174
nigra, Elapsoidea, 290
nigra, Elapsoidea s., 290
nigra, Laeerta, 229, 230
nigra, Pelouiodusa, 174

nigra, Pseudohaje, 293
nigrescens, Rana, 342
nigrescens, Sternotherus, 175
nigricans, Leptotyphlops, 246
nigricans, Phryniscus, 308
nigricans, Testudo, 175
nigricans, Typhlops, 245, 246
nigricollis, Naja, 292
nigricoUis, Naja n., 292
nigricollis, Stellio, 196
nigripes, Tiliqua, 214
nigroalbus, Typhlops, 240
nigrocollaris, Aparallactus, 286
nigrolineatus, Gerrhosaurus, 224
nigrolineatus, Gerrhosaurus n., 224
nigrolineatus, Onychocephalus, 242
nilotica, Laeerta, 234, 235, 236
nilotiea, Rana, 342
niloticus, Crocodylus, 176, 177
niloticus, Trionyx, 171, 172
niloticus, Varanus n., 235
nitida, Eremias, 232
iiitidulus, Hyperolius, 325
nitschei, Atheris, 304
nitschei, Atheris n., 304
nkukae, Arthroleptis x., 351
iioblei, Hyperolius, 328, 331
iioltei, Geoealamus, 239
notabilis, Euprepes, 209
notatum, Mierosoma, 283
notatus, Ilylambates, 316
Notoa, 174
Notopholis, 228
iiototaenia, Coronella, 275
nototaenia, Hemirhagerrhis n., 275
Nubilia, 181
Nucras, 226, 230
nutti, Rana, 339

nyasalandensis, Phrynomerus b., 355
nyassae, Hyperolius, 328, 329
nyassae, Mehelya, 255
nyassae, Rana, 339
nyassae, Simocephalus, 255
oatesii, Thelotornis k., 275

XXIV

BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

obbianus, Pyxicephalus, 344
obscura, Cassina, 320
obsti, Mabuia, 209
obsti, Mabuya q., 208
obtusa, Dipsas, 271
obtusus, Tarbophis, 270
occidentalis, Agama r., 193
Occident alls, Naja n., 292
occidentalis, Eana, 344
occipitalis, Agama, 194
occipitalis, Rana, 337, 344
oceanica, Cephalochelys, 167
ocellata. Echidna, 301
ocellata, Lacerta, 229
ocellata, Pseudocassina, 316
ocellatus. Coluber, 266
ocellatus, Hylambates, 316
ocellatus, Varanus, 235, 236
octophractus, Crocodilus, 177
oculatus, Hyperolius, 325
Odatria, 235

ogoensis, Phrynobatraehus, 347
oldhami, Chlorophis, 260
Oligolepis, 260
olivacea, Chelonia, 168
olivacea, Coronella, 256
olivacea, Emys, 173
olivacea, Lepidochelys o., 168
olivacea, Natriciteres o., 256
olivacea, Pelomedusa s., 173, 174
olivaceus, Boaedon, 250
olivaceus, Holuropholis, 250, 251
olivaceus, Megablabes, 257
Ollotis, 310
Ommateremias, 233
onunatostictus, Hyperolius m., 330
Onychocephalus, 240
Onychochelys, 166
Onychopbis, 240
Onychopus, 181
Oopholis, 176
Opetiodon, 268
OPHIDIA, 239
Ophinectes, 295

Ophioproctes, 237
Ophiopsis, 218
Ophirhijia, 266
Ophthalmidium, 240
OPISTHOCOELA, 307
OPISTHOGLYPHA, 268
opisthopachys, Typhlops, 243
oppellii, Eumeces, 214
orangensis, Pelomedusa g., 174
orbicularis, Agama, 192
orientalis, Amphisbaena, 238
orientalis, Amphisbaenula, 238
orientalis, Gastropyxis, 263
ornata, Braehysaura, 193
ornata, Pelamis, 295
ornata, Rana o., 345
omatissima, Hildebrandtia, 345
oruatissima, Prosymna, 265
ornatissiiiui, Prosymna a., 265
ornatum, Lycophidion, 252
ornatum, Lycophidion c, 252
ornatus, Hyperolius, 326
ornatus, Leiurus, 181
ovnatuH, Maerophis, 267
ornatus, Pyxicephalus, 337, 345
ornatus, Varanus n., 236
orophilus, Af rixalus b., 323
osborni, Osteoblepharon, 178
osborni, Osteoblepharon t., 178
Osilophus, 309
Osteoblepharon, 178
Osteolaemus, 178
Ota^pis, 309
Otolophus, 309
ouba.nghiensis, Rana, 339
oustak'ti, Chamaeleo, 206
oustaleti, Chamaeleon, 206
ovamboeusis, Kassina s., 321
oweni, Aparallactus j., 286
oweni, Chamaeleo, 196
oxiana, Tomyris, 291
oxyrhinus, Sternotherus, 174
OxyrhjTiehus, 309
oxyrhynchus, Psauunophis, 277

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS XXV

oxyrhynchus, RaJia, 340

oxj-rhynchus, Kana o., 340, 341

oxyrhj-nchus, Rappia, 333

oxyrhj-nchus, Khamphiophis o., 277

Oxytropis, 208

Pachybatraehus, 337

Pachydactylus, 191

pachyderma, Hyperolius, 325, 359

pachyderma, Eappia, 325

Pachysaurus, 235

pakenhami, Lygodactylus g., 187, 188

pakenhami, Phrynobatrachus, 347

Paleorana, 338

Palinia, 176

pallida, Letheobia, 240, 244

pallida, Naia n., 292

pallida, Naja n., 292

pallidus, Hydrus p., 296

pallidus, Typhlops, 244

pallidus, Typhlops b., 244

palmarum, Dasypeltis s., 288

palmipes, Eana, 337

palustris, Crocodilus, 176

pammeces, Typhlops, 244

pannonicus, Ablepharus, 218

pantherinus, Bufo, 310

pantherinus, Hyperolius, 326, 359

pantherinus, Mexalixalus, 326

Pantherosaurus, 235

Pappophis, 268

Paracassina, 320

Paragonatodes, 180

Pararthroleptis, 351

pardalis, Chamaeleo, 197

pardalis, Eremias, 232

pardalis, Testudo, 169

Parhoplophryne, 358

parisiensiiim, Chamaeleo, 196

parked, Atractaspis i., 297

parkeri, Bufo, 312

parkeri, Hemidactylus, 184

parkeri, Hyperolius, 332

parkeri, Hyperolius p., 332

parkeri, Leptopelis, 316, 319

parkeri, Phelsuma m., 190

parkeri, Zonurus, 223

parkeriana, Eana a., 338

Parkerophis, 284

Paroedura, 189

parreysii, Elaphe, 2.") 7

parvilobus, Chaniaeleon, 199

parvulus, Phrynobatrachus, 350

Pataeosaurus, 232

pauciscutatus, Crocodylus n., 178

pavo, Echis, 303

Pediophylax, 232

Pedioplanis, 232

Pegaeus, 309

Pelamis, 295

pelaniis, Hydrophis, 295

Pelaniydrus, 295

Pelamys, 295

Pelasgorum, Chelonia, lt)7

Pelias, 300

Pelomedusa, 173

PELOMEDUSIDAE, 173

Pelophylax, 336

Peltastes, 169

I)cltocephalus, Bufo, 309

Peltonia, 169

Peltophryne, 309

Pelusios, 174

pembae, Leptotyphlops e., 247

pembana, Natriciteres o., 256

pembana, NatrLx o., 256

pembana, Eiopa, 213, 216

pembanum, Lygosoma, 216

penangeusis, Ansonia, 310

Pentonyx, 173

peraffinis, Aparallactus, 28n

percivali, Acontias, 222

perdioolor, Eumeces, 21."

Periaspis, 286

Periops, 257

perpalmatus, Phrj-nobatrachus, 34(i

perrotetii, Mabuya, 211

persimilis, Hemidactylus, 186

perspicillata, Lacerta, 229

XXVI

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

peruvianus, Hemidactylus, 181
petersi, Chamaeleon d., 199
petersi, Hyperolius, 333
petersii, Cliiromantis, 315, 316
petersii, Chiromantis p., 315
petersii, On3'chocephalus, 241
petersii, Xenopus, 308
petersii, Xenopus 1., 308
Petia, 218
petropedetoides, Phrynobatrachus,

348
Phaneropis, 218
Phayrea, 278
Phelsuina, 190
Phenax, 229
Philas, 176

pliillipsi, Atractaspis, 297
phillipsii, Psammophis s., 279
Philoehortus, 227
Philodendros, 159, 277
Philodendrus, 278
Pliilothamnus, 259
Phlyctimantis, 320
phocarum, Coronella, 266
Phryne, 309
Phryniscus, 308
I'hrynobatrachus, 346
Phrynoderma, 336
phrynoderma, Hyperolius, 325
Phrynoidis, 309
Phrynomantis, 355
PHRYNOMERIDAE, 355
PHRYNOMERINAE, 355
Phrynomerus, 355
Phrynomorphus, 309
Phrynopsis, 192, 337
Phumaaiola, 197
Phyllodactylus, 189
phyllophis, Siniocephalus, 254
phylofiniens, Amphisbaena, 238
piequotii, Varanus, 235
pictieauda, Agama, 194
picturatus, Hemidactylus, 188
picturatus, Hyperolius, 325

picturatus, Lygodactylus p., 188

pictus, Chiromantis, 316

pietus, Hyperolius, 325

Pilidion, 240

pinguis, Typhlops, 243

PIPIDAE, 307

pitmani, Hyperolius, 335, 359

pitmani, Prosymna, 265

Placovaranus, 235

planieeps, Chamaeleo, 200

planifrons, Euprepes, 211

planifrons, Mabuya, 211

Planodes, 192

platura, Anguis, 295

platuro, Anguis, 295

platurus, Pelamis, 295

platycephala, Rappia, 332

platycephalus, Hemidactylus, 185

I'latyeeps, 257

Platypholis, 191

platyrhinus, Rappia, 329

platyrhynchus, Typhlops, 243

PLEURODIRA, 173

Pleurostrichus, 225

Pleurotuchus, 224

Pleurotychus, 224

plicatus, Hyperolius, 347

plicatus, Phrynobatrachus, 347

Plocederma, 193

plumbeatra, Periaspis, 286

plumbeiceps, Meizodon, 258

plutonis, Boodon 1., 251

Pnoepus, 181

Podarcis, 229

Podorrhoa, 192

poecilonotus, Arthroleptis, 352

poecilopleuro, Ablepharus, 218

Poecilostolus, 303

poensis, Heterolepis, 254, 255

poensis, Mehelya, 255

Pohlia, 337

polyaspis, Chelonia, 168

Polydaedalus, 235, 236

polygrammicus, Typhlops, 239

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS XXVli

polylepis, Atheris, 304
))olylepis, Calamelaps, 281
polylepis, Calamelaps u., 281, 359
polylepis, Dendraspis, 294
polylepis, Dendroaspis p., 294
POLYPEDATIDAE, 314
ponctuc, Agame, 179
porcata, Dermatochelys, 164
porosus, Crocodilus, 176
porrectus, Coelopeltis, 277
poweri, Kassina, 320, 321
poweri, Xenopus, 30S
jjraeomata, Coronella r., 259
praeornata, Dendrophis, 159, 278
l)rasiiia, Gastropholis, 227
proboscideus, Bufo, 309
I'robrevieeps, 356
Proeoela, 308

procterae, Geodipsas, 255, 25'5
procterae, Malacochersus, 170
procterae, Testudo, 170
Proscelotes, 219, 220
Prosymna, 264
proximus, Atheris, 304
prunicolor, Aporarchus, 237
Psammobates, 168, 169
Psammophis, 277, 278
Psammophylax, 159, 275, 276
Psanimorrhoa, 158, 192
Psammosaurus, 234
Psammoscopus 235
Pseudarthroleptis, 351
Pseudaspis, 266
Pseuderemias, 232
pseudocaretta, Chelonia, 166
Pseudocassina, 316
pseudodipsas, Dendrophis, 273
Pseudohaje, 291, 292
pseudomydas, Chelonia, 166
Pseudoprosymna, 265
Pseudotrapelus, 192
Pseudotyphlops, 239
Pseudoxenopus, 337
Pterosaurus, 197

Ptychadena, 337
puccionii, Hemidactylus, 184
puleher, Migiurtinophis, 270
pulcher, Monitor, 236
puleher, Phyllodactylus, 189
pulchra, Mabuya, 211
pulchra, Rana, 345
pulchra, Tornierella, 320
pulchrouiarmoratus, Hyperolius, 329
pulverulenta, Boiga, 269
pulverulenta, Dipsas, 269
pumila, Lacerta, 196, 197
pumilus, Chamaeleon, 196
pumilus, Hemidactylus, 183
punctata, Hyla, 318
punctata, Lacerta, 213, 214
punctata, Odatria, 235
punctata, Teira, 229
punctata, Trachylepis, 208
punctatissimus, Euprepes, 213
punctatissimus, Hyperolius, 328, 330
punctatolineatus, Aparallactus, 287
punctatostriatus, Enicognathus, 256
punctatus, Acontias, 239, 241
punctatus, Dispholidus t., 274
punctatus, Hemidactylus, 183
punctatus, Philothamnus, 262
punctatus, Typhlops p., 241
puncticulata, Eappia, 331
puncticulatus, Hyperolius, 328
puncticulatus, Hyperolius p., 331
punctulata, Eappia, 333
punctulatus, Mochlus, 214, 215
pusilla, Crumenifera, 333
pusillus, Hyperolius, 333
pygmaeus, Arthroleptis, 351
pygmaeus, Chiromantis, 316
pygmaeus, Hyperolius, 323
pyramidium, Echis c, 303
pyramidum, Scythale, 303
pyrrhocephalus, Euprepes, 211
Python, 162, 247, 248
PYTHONINAE, 247
Pyxicephalus, 336

xxvm

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

quadratomaculatus, Hyperolius, 335,

359
quadrifrons, Amphisbaena, 237, 238
quadrilineatum, Boaedon, 251
quadrivittatum, Boaedon, 251
quadrupes, Anguis, 216
qiiattuorseriatus, Ciiemaspis, 180
quattuorseriatus, Gonatodes, 180
quattuorliiieatus, Coluber, 257
quecketti, Eana, 339
quilensis, Chamaeleo d., 199
quinquedigitata, Lygosoma g., 217
quinquelineatus, Lygodactylus p.,

188
quinquetaeniata, Mabuya q., 208
quinquetaeniatus, Seineus, 208
Rachiodon, 288
Eana, 336
Eanaria, 336
EANIDAE, 335
ranoides, Phrynobatraelius, 348
Eanosoma, 337
Eanula, 337
Eaphiodoii, 288
Rappia, 324
redi. Coluber f ., 300
redii, Vipera, 300
regalis, Dipsas, 269
Eegenia, 235

regularis, Afrixalus d., 322
regularis, Bufo, 310
regularis, Bufo r., 310, 311
regularis, Mcizodon, 258, 259
reichei, Arthroleptis, 351
remivaga, Caretta, 168
Rena, 246

renschi, Hyperolius, 328
REPTILIA, 163
resimus, Causus, 298, 299
resimus, Heterophis, 298
reticulata, Boa, 248
reticulata, Dendroechis, 293
reticulatus, Dipsadomorphus, 271
retieulatus, Eumeces, 215

reuteri, Typhlops, 245
revoili, Eremias, 231
revoili Latastia 1., 231
RHACOPHORIDAE, 314, 355
Rhaebo, 309
Rhanmophis, 263, 264
Rhaniphiophis, 276
Rhampholeon, 197, 206
Rhamphophis, 279
Rhamphostoma, 246
Rhamphotyphlops, 239
Rhinaspis, 300
Rhinechis, 300
Rhinoealamus, 281, 282
rhinoceratus, Chamealeo, 197
rhinoceros, Bitis g., 302
Rhinoptyon, 235
Rhinoscincus, 220
Rhinotyphlops, 239
RHIPTOGLOSSA, 196
rhodesianus, Cordylus c, 223
rhodesianus, Pelusios n., 175
rhodoscelis, Hyperolius, 326, 328,

329
rliomlieata, Sepedon, 298
rhombeatus, Causus, 298
rhombeatus, Coluber, 159, 275, 276
rhombeatus, Trimerorhinus, 276
rhonibifer, Croeodilus, 176
richardsonii, Valernesia, 181
Riopa, 213, 216
robecchii, Brookesia, 206
robecchii, Rhampholeon, 206
robustus, Croeodilus, 177
robustus, Pachybatrachus, 337
rogersi, Hoplophrj-ne, 358
rohrbeoki, Mabuia m., 209
rondoensis, Amphisbaena, 238
rondoensis, Bufo m., 313
rondoensis, Melanoseps a., 221
rondoensis, Typhlops t., 243
roperi, Chamaeleo d., 199
roperi, Chamaeleou, 199
rosei, Arthroleptis, 345

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS XXLS

lossii, Rappia, 325
rostrata, Argyrogena, 258
rostrata, Atractaspis, 29S
rostrata, Atractaspis b., 298
rostrata, Caretta, 166
rostrata, Hypogeophis, 305
rostratus, Causus, 299
rostratus, Rlianiphiophis, 276, 277
rostratus, Ehampliiophis o., 277
Rothsehiklia, 320
rothschildi, Mabuia, 211
rothsehildi, Thrasops, 264
rouclieti, Aparallactus n., 286
Rouleophis, 285
rovumae, Hyperolius p., 333
rouxi, Arthroleptis, 350
rouxi, Phrynobatraehus, 350, 359
rubripes, Hyperolius, 332
rubropunetata, Dipsiua, 277
rubropunctata, Lacerta, 232
rubropuuctatus, Rhamphiophis, 277
rudicollis, Uarauus, 235
rudis, Chamaeleo b., 201
rudis, Chamaeleon, 201
rudolfensis, Philochortus i.. 228, 359
rudolfianus, Trionyx t., 172
rufescens, Chiromantis, 315
rufescens. Coluber, 272
rufescens, Coronella, 271
rufescens, Lacerta, 21-4
rufescens, Polypedates, 315
rufula, Coronella, 2'49
rufulus. Coluber, 249, 250
rufulus, Tragops, 274
rufus, Leptopelis, 319
rugegensis, Leptopelis, 319
rugiceps, Eremias, 233
rugosa, Pseudocassina, 316
rugosus, Cliiromantis, 316
rungweensis, Atheris n., 304
rungwensis, Arthroleptis, 349
rungwensis, Phrynobatraehus, 349
rungwensis, Probreviceps m., 357
ruppelli, Agama r., 193

ruppellii, Caouana, 168
ruspolii, Asthenophis, 265
ruspolii, Chamaeleo d., 199
ruspolii, Hemidactylus, 185
russelli, Coluber, 300
russellii, TortrLx, 244
Sabrina, 246
Saiphos, 216

sakalava, Hemidactylus, 185
SALIENTIA, 307
salvator, Stellio, 235
samharica, Lacerta, 231
sanctijohannis, Paroedura, 189
sanguinolenta, Lacerta, 192
sansibarica, Rappia, 332
sansibaricus Hyjierolius c, 332
sansibaricus, Philothanmus p., 262
Sarea, 237
SAURIA, 178
Sauroceras, 197
Saurophis, 225
saurus, Stellio, 236
savignyi, Euprepes, 208
savignyi, Rana, 342
savignyi, Trapelus, 179
savorgnani, Heterolepis, 254
savorgnani, Mehelya c, 254
saxatilis, Stellio, 192
scaber. Coluber, 288
scaber, Dasypeltis s., 288
scabra, Dasypeltis, 288
scabrieeps, Teratolepis, 182
Scalabotes, 186
Scaphiophis, 264
Scelarcis, 229
Scelotes, 219, 220, 221
sr-hebeni, Arthroleptis, 354
schefifleri, Arthroleptis, 349
scheffleri, Coronella, 259
seheffleri, Hyperolius, 329
scheffleri, Lygodactylus, 186
scheffleri, Lygodactylus f., 186
schereri, Ranosoma, 337, 344
Schismaderma, 309

XXX

BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Schistonietopuni, 305
sclilegeli, Leionotus, 267
schlegeli, Lejonotus, 267
schlegelii, Spliargis c, 164
schlegelii, Typhlops s., 241
schmidti, Miodon g., 283
schmidti, Thrasops j., 264
schneideri, Megalixalus, 322
achneideri, Pelamis, 295
schoenebecki, Arthroleptis, 352
vschoensis, Kinixys, 170
schopfiij Testudo, 171
Schoutedenella, 350, 351
sehoutedeni, Atraetaspis, 297
schoutedeni, Chamaeleo b., 200
sehoutedeni, Megalixalus, 323
schubotzi, Arthroleptis, 351
schubotzi, Chamaeleo b., 201, 359
schubotzi, Chaniaeleon b., 201
schubotzi, Chlorophis, 262
schubotzi, Hyperolius, 326
schubotzi, Mabuia m., 209
schubotzi, Rana, 341
SCINCIDAE, 208
Scincodipus, 220
Scoleconiorphus, 306
8colecoseps, 222
Scoliophis, 257

scriptus, Hyperolius, 330, 335, 359
scutifrons, Leptotyphlops, 246
Scutiger, 308
sebae. Coluber, 248
sebae, Euprepes, 208
sebae. Python, 248
somiannulatus, Telescopus, 270
semiannulatus, Telescopus s., 270
semiannulis, Lycophidion, 253
semiornatn, Coronella, 259
seniiornatus, Meizodon s., 258
seniivariegata, Dendrophis, 262
semivariegatus, Philothamnus s., 259,

262
Semnodactylus, 321
senegalensis, Chamaeleo, 197

senegalensis, Chamaeleo s., 197

senegalensis, Cystignathus, 320, 321

senegalensis, Kassina, 321

senegalensis. Monitor e., 236

Sepacontias, 214

sepiformis, Scincus, 225

Sepomorphus, 220

seps, Lacerta, 225

Sepsina, 219, 220

septemlineatus, Lygodactylus p., 188

septemstriatus, Tj'phlops, 239, 246

septentrionalis, Agama r., 193

serpens, Lacerta, 216

SERPENTES, 239

severa, Vipera, 302

sextaeniata, Eremias, 233

sextaeniata, Eremias s., 233

seychellensis, Megalixalus, 321

seychellensis, Sternothaerus n., 175

Seytina, 164

shirana, Ahaetulla, 261

shirana, Duberria 1., 267

shiranum, Homalosonia, 267

Shrevea, 237, 238

Siagonodon, 246

Siaphos, 217

sibilans, Coluber, 278, 279

sibilans, Psammophis s., 278, 279

sibiricuni, Chorisodon, 278

Riebenrockia, 254

signata, Leptotyphlops, 246

signifer, L^ptopelis, 317, 319

Silurana, 307

silurophaga, Grayia, 267

Simalia, 247

Simocephalus, 254

simoni, Chamaeleo, 199

simplex, Tetradactylus f., 225

siuaita, Agama, 192, 193

sinuata, Pelamis b., 295

sinuatus, Pelusios, 174, 175

sinuatus, Sternotherus, 175

sjostedti, Dendraspis, 294

sloanii, Scincus, 208

LOVERIDGE: E. AFRICAN REPTILES AND AMPHIBIANS XXxi

smaragdina, Dendrophis, 263
smaragdiiia, Gastropyxis, 263
smaragdinus, Hyperolius k., 332
smithi, Co]u))er f., 258
smithi, Eieniias, 233
smithi, Zamenis, 258
smithii, Philothamnus, 262
smythii, Grayia, 2(37
somalica, Bitis a., 302
somalica, Bitis 1., 302
somalica, Meizodon, 25S
somalica, Rana, 338
somalica, Riopa m., 215
somalicus, Anops, 237
somalieus, Lygodactylus, 186
somalicus, Lygodactylus s., 186
somalicuK, Tarbopliis d., 270, 359
somalicus, Telescopus d., 270
somaliensis, Mehelya, 254
Spalerosophis, 257
speciosus, Coluber, 248
spectrum, Chamaeleo, 206
spekii, Eremias, 233
spekii, Eremias s., 233
spekii, Kinixys, 170
Spelaeophryne, 356
sphaeropholis, Chamaeleon, 198
Sphaerotheca, 337
Sphargis, 164
Sphenorhina, 222
Sphenosoma, 214
spinidactyla, Rana, 342
spinifroiis, Hyperolius, 322
spinosus, Bufo, 310
spinosus, Chamaeleo, 202
spinosus, Chamaeleon, 202
spiuosus, Megalixalus, 321
Spondylurus, 208
SQUAMATA, 178
squama ta, Eretmochelys, 166
Squamieilia, 214
squamiger, Atheris, 303, 304
squamigera, Echis, 304
squamosa, Caretta, 166

squamulatus, Hemidactylus, 182, 183
squamulatus, Hemidactylus a., 182
squamulosa, Ichnotropis, 234
stcindachnerii, Bufo, 312
stellata, Testudo, 169
stellatus, Hemidactylus, 184
stellatus, Pachydactylus b., 191
stellatus, Tupinambis, 236
Stellio, 192

stelzneri, Atelopus, 308
stenocephala, Rana, 343
Htenodactylus, 179
st«nodactylus Arthroleptis s., 352,

353
stenodactylus, Ascalabotes, 179
stenophthalmus, Heterolepis, 255
stenophthalmus, Mehelya, 255
.Stenorhabdium, 265
Stenorhyuchus, 346
Stenostoma, 245, 246
Stenostomophis, 246
fiternothaerus, 175
Sternotherus, 175
stevensoni, Lygodactylus, 187
sthonodactylus, Ascalabotes, 179
sthenodactylus, Stenodactylus s., 179
Rtigmochelys, 168, 169
stormsi, Boulengcrina, 291
stormsi, Boulengerina a., 291
striata, Eremias, 233
striata, Euprepis, 214
striata, Mabuya s., 213
striatum, Tropidolepisma, 213
striatus, Eremias b., 233
striatus, Typhlops, 239
.strigata, Sphaerotheca, 337
strigatus, Lygodactylus, 187
striolatus, Hyperolius, 329, 330
Strongylopus, 336
stuhlmanni, Hyperolius, 326
stuhlmanni, Ligonirostra, 265
stuhlmanni, Prosymna a., 265
subfasciatus, Platyceps, 257
subfasciatus, Rachiodon, 289

XXXll

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

subfulva, Naja m., 293
sublaevis, Boltalia, 181
subnig-er, Pelusios, 174
pubniger, Stemothaerus, 174
subnigra, Testudo, 174, 175
subobscurus, Hydrus p., 296
subocularis, Atheris, 304
subrufa, Pelomedusa s., 173
subrufa, Testudo, 173
subsigillata, Rana, 337
substriatus, Hyperolius, 330
substriatus, Hyperolius p., 330
subtaeniatus, Psammophis s., 280
succinea, Nucras, 230
suohus, Crocodihis, 177
sudanesis, Kakophrynus, 354
sudaneiisis, Psammophis s., 280
sulcata, Testudo, 169
simderallii, Eumices, 215
sunderallii, Euprepes, 213
sundevallii, Eumeces, 214
sundevallii, Riopa s., 213, 215
sundevallii, Tcnmorhynchus, 265
sundewallii, Temiioihynehus, 265
superciliaris, Chamaeleo, 206
superciliaris, Rana o., 341
superciliaris, Vipera, 301
swynnertoui, Chirindia, 237
sykesii, Barycephalus, 193
sylvatiea, Sehoutedenella, 351
symetrica, Rappia, 329
SYNAPTOSAURIA, 178
Systoma, 357
Tachybates, 181
Tachygia, 214
Tachyscelis, 232
taeniata, Causus r., 299
taeniatus, Amplorhinus, 275
taitanus, Boulengerula, 306
taitanus, Bufo, 313
taitanus, Bufo t., 313
taitanus, Euprepes, 211
taitanus, Ileniisus, 354
taitensis, Chamaeleo, 204

tanae, Riopa, 213, 214
tancredi, Ablepharus, 219
tanganicana, lehnotropis, 234, 359
tanganicus, Hemidactylus, 185
tanganicus, Psammophis b., 280
tanganus, Leptopelis, 319
tanganyikae, Chilorhinophis g., 284
Tanieremias, 233
Tanoa, 174
Tapaya, 192
Tapelus, 192
Taphriomantis, 316
Taphrometopon, 278
tarapacona, Thalassochelys, 167
Tarbophis, 270
tasmani, Hemidactylus, 186
taurica, Laecrta, 229
tavetanus, Chamaeleo f., 203
tavetanus, Cliamaeleon, 203
tavetensis, Oiamaeleon, 203
taylori, Hemidactylus, 182
taylori, Teratolepis, 182
tchadensis, Zamenis, 259
Tectovaranus, 235
Teira, 229
Telcovipera, 300
Telescopium, 270
Telescopus, 270, 271
Temnorhynchus, 264
tempeli, Chamaeleo, 202
tempeli, Chamaeleon, 202
temporale, Stenorhabdium, 265
temporalis, Brookesia, 207
temporalis, Chamaeleon, 207
temporalis, Psammophis b., 278
temporaria, Rana, 336, 337
Temsacus, 176
tenue, Ophthalmidium, 244
tenuior, Chamaesaura, 226
tenuior, Chamaesaura a., 226
tenuis, Chamaeleo, 203
tenuis, Chamaeleon, 203
tenuis, Chelonia, 166
Teratolepis, 182

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS XXXlll

tessellata, Mabuya, 211

tcssellatiini, Typhlops, 246

TESTUDINATA, 163

Testudiuella, 169

TESTI^DINES, 163

TESTUDINIDAE. 168

Testiulo, 163, 168

tetensis, Pachydactylus, 191

Tetracheilostoma, 246

tetradactyla, Sepsina, 220, 221

Tetradactylus, 225

tetradactylus, Chalcides, 225

tetradactylus, Seelotes t., 219, 221

teti-aspis, Osteolaemus, 178

tetraspis, Osteolaemus t., 178

tettensis, Hyperolius, 332

tettensis, Onychoeephalus, 243

tettensis, Psammophis s., 279

tettensis, Typhlops t., 243

thabanchueusis, Semnodactylus, 321

Thalassochelys, 167

Thalestris, 229

THECOPHORA, 163

theileri, E^na, 341

Theleus, 250

Thelotornis, 274

Thermophilus, 159, 233

Thetia, 229

tliolloni, Grayia, 267

thomasi, Lygosoma, 216

thomasi, Psammophis, 279

thomensis, Scalabotes, 186

Thrasops, 263

thunbergii, Caretta, 165

Thyrus, 219, 220

tibatieusis, Chamaeleon s., 198

tigrina, Mesocoronis, 300

tigris, Chamaeleo, 197

tigris, Python, 248

Timon, 229

timoremsis. Monitor, 235

togoensis, Hemidactylus b., 184

togoensis, Rana, 345

Tolarenta, 179

Tomopterna, 337
Tomyris, 291
Tornierella, 320
tornieri, Chamaeleon, 204
tornieri, Chamaeleon b., 200
tornieri, Crotaphopeltis h., 271
tornieri, Hemidactylus, 183
tornieri, Hyperolius, 335, 359
tornieri, Leptodira, 271
toinieri, Malaeochersus, 169
tornieri, Neetophryne, 314
tornieri, Neetophrynoides, 314
tornieri, Testudo, 169, 170
tornieri, Typhlops, 242
Toxieoa, 303
Toxicodryas, 268
Toxodon, 257
Trachylepis, 208
translueens, Hyperolius, 334
transvaalensis, Psammophis, 280
Trapeloidis, 192
Trapelus, 192
Trapetus, 192
triangularis, Coronella, 267
Triceras, 196
Tricheilostoma, 246
tricolor, Pelamis p., 296
triedrus, Elasmodactylus, 192
Triglyphodon, 268
trigonatus, Coluber, 268
Trigonophis, 270
trilamina, Echidnoides, 300
Trimerorhinus, 276
triuodis, Rana, 341
Trioceros, 196
TRIONYCHIDAE, 171
Trionyx, 171
tristis, Monitor, 235
tritaeniata, Coronella, 276
tritaeniata, Rhagerrhis, 276
tritaeniatus, Psanimophylax t., 276
tritaeniatus, Rhagerrhis, 276
tritaeniatus, Trimerorhinus, 276
Tritonopsis, 230

XXXIV

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

triunguis, Testudo, 171, 172
triunguis, Tiionyx, 172
trivirgatus, Psammophis p., 279
trivittatus, Scincus, 220
tropicalis, Silurana, 307
tropidolepis, Hemidactylus, 182
Tropidopholis, 228
tropidopholis, Bedriagaia, 227
tropidosteniuni, Cordylus c, 223
tropidosternum, Zonurus, 223
truncatus, Argyrophis, 244
Trypheropsis, 337
tuberculata, Agania, 193
tuberculata, Testudo, 164
tuberculosa, Pyxieeplialus, 345
tuberculosa, Rana, 345
tuberculosus, Elasmodactylus, 191
tuberculosus, fleeko, 160, 185
tuberculosus, Hemidactylus, 181
tuberculosus, Paehydactylus, 191
tuberculosus, Phyllodactylus, 189
tuberilinguis, Hyiterolius, 332
tuberilinguis, Hyperolius c, 332
tumbensis, Psammophis s., 278
tumboensis, Dipsas g., 269
tupinambis, Lacertus, 236
turcica, Boa, 248
turneri, Aparallactus, 287
turneri, Homodactylus, 191
turneri, Paehydactylus b., 191
turuensis, Agaraa a., 195
Tylanthera, 257
Typhlina, 239
Typhlinalis, 240
TYPHLOPIDAE, 239
Typhlops, 239
typhonia, Rana, 309
typhonius, Osilophus, 309
typicus, Pleurotuchus, 224
typus, Anodon, 288
typus, Bucephalus, 273
typus, Dispholidus, 273
Tyrse, 171
Tzarevscya, 300

Uaranus, 234

ubangensis, Aparallactus, 285
udjidjiensis, Hyperolius, 329
uelensis, Atractaspis i., 297
ufipae, Agama a., 195
ukamiensis, Arthroleptis, 353
ukerewensis, Lj'godaetylus p., 188
ukerewensis, Nucras, 230
ukingensis, Arthroleptis, 350
ukingensis, Cordylus, 223
ukingensis, Phrynobatrachus u., 350
ukingensis, Zonurus, 223
uluguruensis, Afrixalus, 323
uluguruensis, Aparallactus, 287
uluguruensis, Arthroleptis s., 350,

352, 353
uluguruensis, Boulengerula, 306
uluguruensis, C^aniaeleo f., 204
uluguruensis, Hoplophryne, 358
uluguruensis, Lcptopelis, 319
uluguruensis, Megalixalus, 323
uluguruensis, Natriciteres o., 256
uluguruensis, Natrix o., 256
uluguruensis, Probreviceps, 357
uluguruensis, Scelotes, 219, 220
uluguruensis, Scoleeomorphus k., 307
uluguruensis, Typhlops, 244
umbelluzianus, CTiiromantis, 315
undata, Lacerta, 232
undecinistriatus, Typhlops, 246
undulata, Rappia, 329
undulatus, Hyperolius, 328
unguieulata, Rhagerrhis, 277
unieolor, Bdellophis, 305
unic'olor, Boaedou, 250
unic-olor, Calamaria, 281
unieolor, Calamelaps u., 281
unieolor, Dipsadoboa, 269
unieolor, Guyomarehia, 285, 286
unieolor, Hyperolius, 323
unieolor, Lycodon, 251
unieolor, Megalixalus f., 322
unieolor, Miodon, 283
unieolor, Rachiodon, 289

LOVERIDGE : E. AFRICAN REPTILES AND AMPHIBIANS XXXV

unk'olor, Simoeephalus, 254
iinidecimstriatus, Tvphlops, 246
iinitaeniatus, Typlilops, 242
unitaeniatus, Tvphlops u., 242
upembae, Megalixalus 1., 232
Uperolia, 324
Uraeus, 291
Uriechis, 285
Urobelus, 283
iirsinii, Pelias, 300
urunguensis, Bufo, 312
usambarae, Agama a., 194
usambarae, Agama c, 194
usambarae, Leptopelis, 319
usambarae, Phrynopsis, 344
usambarica, Parhoplopyhryne, 358
usambarieus, Breviceps, 357
usambaricus, Parhoplophryne, 358
usaramoae, Hyperolius, 334
ushoranus, Bufo, 312, 314
uthmoUeri, Chamaeleo, 203
uthmolleri, Chamaeleo f., 203, 359
uzuiigueusis, Bufo t., 313
uzungwense, Lyeophidion c, 252
uzungwensis, Lyeophidion c., 252
uzungwensis, Melanoseps a., 222
uzungwensis, Rana ni., 342, 343
vagus, Arthroleptis, 353
vaillanti, Agama, 193
vaillanti, Elaposchema, 282
Valernesia, 181
valida, Dipsas, 268
vanrooyeni, Phrynobatrachus, 350
VARAXIDAE, 234
Varanus, 234
varia, Echis, 303
varia, Laeerta, 234
varia, Mabuya v., 212
variabile, Agame, 192
variabilis, Arthroleptis, 353
variabilis, Hyperolius, 327, 328, 330
variabilis, Prosymna, 265
variabilis, Psammophylax, 276
variabilis, Psammophylax t., 276

variegata, Nannophryne, 310
variegata, Pelamis b., 29')
variegatum, Alopecion, 251
variegatus, Euprepes, 213
variegatus, Heleionomus, 248
variegatus, Hemidactylus, 188
varius, Euprepes, 212
vassei, Prosymna, 265
vaurescecae, Chamaeleon j., 205
vauereselli, Algyroides, 229
vauereselli, Laeerta, 229
vauerocegae, Geodipsas, 255
velox, Laeerta, 232
ventromaculatus. Coluber, 257, 258
venusta, Rana, 342
venusta, Rana m., 342
vermicularis, Hyperolius, 330
vermiculata, Rappia, 330
vermiculatum, Engystonia, 355
vermiculatus, Hylambates, 319
vermieulatus, Leptopelis, 317, 319
vernayi, ChJorophis, 261
vernayi, R^ana, 341
verrucosus, Hylambates, 320, 359
versicolor, Phrynobatrachus, 347, 348
verus, Cordylus, 223
vibicaria, Levirana, 337
victor ianus, Xenopus, 307
victorianus, Xenopus 1., 307, 308
vinekei, Cliamaeleo, 198
violacea, Oxybelis, 274
Vipera, 300
VIPERIDAE, 296
virgata, Chelonia, 165
virgata, Coelopeltis, 250
virgatus, Boaedon f., 251
virgiuica, Coronella, 272
viridiflavus, Coluber, 257
viridiflavus, Eucuemis, 324
viridiflavus, Hyperolius, 325, 328
viridiflavus, Hyperolius v., 324
viridiflavus, Typhlops, 241
viridis, Anoplodipsas, 269
viridis, Bucephalus, 273

XXXVl

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

viridis, Leptophis, 293
viridis, Naja h., 291
viridis, Testudo, 165
viridis-squamosa, Testudo, 165
viridiventris, Dipsadoboa u., 269
vittata, Gastropholis, 226
vittatus, Bdellophis, 306, 307
vittatus, Bufo, 312, 314
vittatus, Hemidactylus, 183
vittatus, Scoleeomorphus, 306
vittiger, Hyperolius, 323
vivax, Coluber, 270
vivax, Tarbophis, 270
vivipara, Lacerta, 229
vivipara, Nectophrynoides, 314
vivipara, Pseudophryne, 314
v-nigrum, Naja, 298
vosmaerii, Hagria, 213
vosmaerii, Scincus, 213
vosseleri, Chamaeleon f., 204
vossii, Boodon, 253
vulgaris, Bufo, 309
vulgaris. Cerastes, 300
vulgaris, Crocodilus, 177
vulgaris, Testudo, 165
vulgaris, Vipera, 300
wahlbergii, Ablepharus, 218, 219
walilbergii, Arthroleptis, 350
wahlbergii, Cryptoblepharus, 219
wahlbergii, Euprepes, 213
walbergi. Gecko, 190
warreni, Calanielaps, 281
warreni, Calamelaps u., 281
wauensis, Mabuia m., 209
wealii, Cassina, 321
weberi, Lygosoma, 217
werneri, Aparallactus, 284, 287
werneri, Chamaeleo, 205
werneri, Chamaeleon, 205
werneri, Chamaeleon f., 204
werneri, Crotaphopeltis, 271, 359
werneri, Cynodontophis, 283
werneri, Hemidactylus, 183
werneri, Leptodira, 271

werneri, Phrynobatrachus w., 346

werthi, Nectophryne, 314

wettsteini, Hyperolius, 327, 328

wettsteini, Pelomedusa s., 174

whytii, Arthroleptis, 351, 353

whytii, Arthroleptis s., 352

whytii, Glypholycus, 250

whytii, Lycodonomorphus r., 250

wiedholzi, Afrixalus b., 323

wilkinsonii, Tolarenta, 179

williamsi, Lygodactylus p., 189

wilsoni, Ablepharus, 219

wittei, Kassinula, 321

wittei, Phrynobatrachus, 339

wittei, Rana, 339

wolffi, Chamaeleon t., 202

wolterstorffi, Diplodactylus, 189

wolterstorffi, Phyllodactylus, 189

woosnami, Atheris, 304

worthingtoni, Bitis, 301

worthingtoni, Crocodylus n., 177

Xenagama, 193

xenodactylus, Arthroleptis, 351

XENOPIDAE, 307

Xenopus, 307

xenorhinus, Chamaeleo, 203

xenorhinus, Chamaeleon, 203

Xenurophis, 267

xerampelina, Chiromantis, 315, 316

Xystrolepis, 208

yakoma, Naia, 293

youngi, Kinixys, 171

Zamenis, 257

zanzibaricus, Gerrhosaurus, 224

zavattari, Stenodactylus s., 179

zechi, Gerrhosaurus m., 224

zenkeri, Rana, 338

zombensis, Kinixys b., 170

Zonurus, 223

Zootoca, 229

zuluensis, Kinixys b., 170

zuluensis, Pelusios s., 17;")

Zygnis, 219, 237

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 117, No. 3

THE SPIDER GENERA CRVSTULINA AND STEATODA

IN NORTH AMERICA, CENTRAL AMERICA, AND

THE WEST INDIES (ARANEAE,

THERIDIIDAE)

By Herbert W. Levi

CAMBRIDGE, MASS., U. S. A.

printed for the museum
August, 1957

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

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Bulletin of the Museiun of Comparative Zoology

AT HARVARD COLLEGE

Vol. 117, No. 3

THE SPIDER GENERA CBUSTULINA AND 8TEAT0DA

IN NORTH AMERICA, CENTRAL AMERICA, AND

THE WEST INDIES (ARANEAE,

THERIDIIDAE)

By Herbert W. Levi

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

August, 1957

No. 3 — The Spider Genera Crustulina and Steatoda

in North America, Central America,

and the West Indies (Araneae,

Theridiidae) .

By Herbert W. Levi

This paper is one of the last in a series of generic revisions
in the family Theridiidae of North America, Central America
and the West Indies.

The genera revised in this paper are characterized by having
a large colulus between the anterior spinnerets, lacking teeth on
the posterior margin of the chelicerae, and having the lateral
eyes close to each other or touching.

The spiders here placed in Steatoda have previously been dis-
tributed among four genera : Asagena, Lithyphantes, Steatoda
and Teutana. The characters which have been used to separate
the four genera are no longer useful as they are based on the
limited European fauna. The Steatoda of previous authors was
characterized by having the anterior median eyes larger than
the others; however, several species placed in Lithyphantes may
have individuals with anterior median eyes the largest (^S".
medialis; S. pulcher). Lithyphantes has been characterized as
having the lateral eyes slightly separated from each other;
however, this character is also present in Steatoda, while in some
species of Lithyphantes, the lateral eyes touch. The few species
of Teutana have the eyes subequal in size and the lateral eyes
touching each other, a combination of characters hardly dif-
ferent from that found in many Lithyphantes species. Several
species placed in Lithyphantes had their opposite sexes described
in Asagena. The shape of the labium and the shape of the
sternum are unreliable characters. The latter depends in part
on the size of the spider ; the former is a characteristic of indi-
vidual species. The carapace is narrow and long in some, usually
the smaller, species ; almost as wide as long in the largest. In
several species of Lithyphantes, the paracymbial hook is on the
mesal side in the cymbium of the male palpus; in others placed
in Lithyphantes, and in species previously placed in Steatoda
and Teutana, it is on the ectal side. However, no sharp line

368 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

can be drawn here either ; there are species which are inter-
mediate.

The idea that some of these genera ought to be combined is not
new. Emerton placed species of Teutana in Steatoda. F.O.P.
Cambridge (1902, Biologia Centrali Americana, Araneidea, vol.
2, p. 374) wrote that "The spiders referred to this genus
[Teutana] are very difficult to separate by any reliable character
from those included under Lithyphantes, though as far as one
can judge, from the material at hand, the contiguity of the lat-
eral eyes may prove constant." We now know that this latter
character did not prove constant. Other authors also have com-
mented on the difficulty of separating these genera.

This revision is based on material in the Museum of Com-
parative Zoology. Dr. W.J. Gertsch has made available to me
the large collections of the American Museum of Natural His-
tory, for which I would like to thank him. Other collections
were made available by Dr. A.M. Chickering; Mrs. D, Frizzell
(Dr. H. Exline) ; Mr. V. Roth; Mr. R.X. Schick; and Dr. R.V.
Chamberlin sent specimens of the Steatoda hipunctata group.
Mr. Schick has raised male Steatoda palomara to maturity,
making it possible to figure its palpus for the first time. Mr.
E. Browning of the British Museum (Natural History) sent a
syntype of Steatoda americana O.P. Cambridge, and a palpus
of the type of Lithyphantes quaesita, as well as other specimens.
Dr. H. Wiehle of Dessau has sent some male Steatoda castanea
from Germany. I wish to express my sincere thanks for the
cooperation of these colleagues.

The bibliographies of holarctic and cosmopolitan species are
not complete ; if needed, they can be consulted in Roewer's recent
Katalog or in Bonnet's Bibliographia.

The drawings of palpi illustrate the left ones. Although keys
are usually not necessary to determine spiders to species, two
keys are included to point out differences in genitalia of the
two species groups of Steatoda.

Cbustulina Menge

CrustuUna Menge, 1868, Schr. Naturf. Gesell. Danzig, new ser., vol. 2, p.
168. Type species: CrustuUna guttata (Wider).
Description. Small theridiid spiders (1-3 mm. total length).

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 369

Carapace longer than wide. Carapace and sternum covered by
large elongate dark tubercles, which are the bases of fine hairs.
Anterior eye row procurved, posterior slightly recurved. Eyes
subequal in size. Anterior median eyes their diameter apart,
almost touching laterals. Posterior medians one diameter apart,
one diameter or slightly less from laterals. Height of clypeus
equals three to four diameters of anterior medians in female,
four to five diameters in male. Chelicerae with one large tooth
on anterior margin, posterior margin without teeth. Legs rela-
tively short, first or fourth legs longest, third shortest. A tarsal
comb present on fourth tarsus. Abdomen subspherical, slightly
flattened dorso-ventrally. Colulus large. Sclerotized ridge on
anterior end of abdomen of males, as in Steatoda, but this struc-
ture also present in females.

Genitalia. Epigynum illustrated by Figures 2 and 5. Palpus
with usual structures (Fig. 9) ; however, cymbium has a mesal
process which may be homologous to the paracymbial hook in
other genera.

Diagnosis. This genus differs from most theridiid genera by
having a large colulus, and lacking teeth on the posterior
margin of the chelicerae ; it differs from Steatoda in having large
dumb-bell shaped tubercles on carapace and sternum, in having
a sclerotized ring around the pedicel on the abdomen of females
and in having a median distal process on the cymbium of the
male palpus.

Remarks. Since many species placed in Crustuli7ia probably
do not belong to it, its distribution is difficult to judge. Crustu-
lina lascivula Keyserling, (1886, Die Spinnen Amerikas, vol. 2,
pt. 2, p. 39, pi. 12, fig. 155, 9 ), is Paidisca unimaculata (Emer-
ton).

Crustulina guttata (Wider)

Thcridioji guttatum Wider, 1834, Abhandl. Mus. Senckenberg, vol. 1, p. 241,

pi. 16, fig. 7, $ , 5 .
Crustulina (juttata, Wiehle, 1937, in Dahl, Die Tierwelt Deutschlands, pt. 33,

p. 191, figs. 194-199, $,<i. Loeket and Millidge, 1953, British spiders,

vol. 2, p. .31, figs. L'Od, 35a,b,e, 9 , <5 .

This species has not been found in America. The many
American references to the name Crustulina guttata are actuallj'

370

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

either C. altera or C. stricta. Both are very similar to it, aud
only details of the genitalia distinguish C. guttata.

Crustulina sticta (O.P. Cambridge)
Figures 1-3, 7 ; Map 1.

Theridion stictum O.P. Cambridge, 1861, Aim. Mag. Nat. Hist., ser. 3, vol.

7, p. 432. Emerton, 1876, Psyche, vol. 1, p. 129; 1877, Proc. Boston

Soc. Nat. Hist., vol. 19, p. 70.
Steatoda sticta, Thorell, 1873, Remarks on synonyms of European spiders,

p. 439.
Crustulina sticta, Simon, 1881, Les Araelinides de France, vol. 5, p. 158.

Banks, 1895, Ann. New York Acad. Sci., vol. 8, p. 417; 1910, Bull.

U.S. Natl. Mus., no. 72, p. 21. Petrunkevitch, 1911, Bull. Amer. Mus.

C. sticto

Map 1. American distribution of Crustulina sticta (O.P. Cambridge).

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 371

Nat. Hist., vol. 29, p. 172. Crosby and Bishop, 1928, Mem. Cornell
Agr. Exp. Sta., no. 101, p. 1039. Emerton, 1930, Publ. Nantucket
Maria Mitchell Assoc, vol. 3, p. 164. Eoewer, 1942, Katalog der
Araneae, vol. 1, p. 399. Kaston, 1948, Bull. Connecticut Geol. Nat. Hist.
Surv., no. 70, p. 75, figs. 34, 2120,9,5. Locket and Millidge, 1953,
British spiders, vol. 2 p. 53, figs. 35d,e,f, 9 ,$ . Kaston, 1955, Trans.
Illinois Acad. Sci., vol. 45, p. 166. Bonnet, 1955, Bibliographia
Araneorum, vol. 2, p. 1256.
Crustulina borealis Banks, 1900, Canadian Ent., vol. 32, p. 98; 1910, Bull.
U.S. Natl. Mus., no. 72, p. 21. Petrunkevitch, 1911, Bull. Amer. Mus.
Nat. Hist., vol. 29, p. 172. Emerton, 1920, Trans. Roy. Canadian Inst.,
vol. 12, p. 312. Worley, 1932, Publ. Univ. Washington Biol., vol. 1,
p. 23. Eoewer, 1942, Katalog der Araneae, vol. 1, p. 399; Gertseh, 1946,
in Procter, Biol. Surv. Mt. Desert Reg., pt. 7, p. 519. Chamberlin and
Ivie, 1947, Bull. Univ. Utah, biol. ser., vol. 10, no. 3, p. 25. Hackman,
1954, Acta Zool. Fennica, vol. 79, p. 48. Bonnet, 1955, Bibliographia
Araneorum, vol. 2, p. 1252.
Crustulina palUpes Banks, 1906, Proc. Ent. Soc. Washington, vol. 7, p. 96,
pi. 2, fig. 15, 9 ; 1910, Bull. U.S. Natl. Mus., no. 72, p. 21. Petrunke-
vitch, 1911, Bull. Amer. Mus. Nat. Hist., vol. 29, p. 172. Worley, 1932,
Publ. Univ. Washington Biol., vol. 1, p. 24. Eoewer, 1942, Katalog der
Araneae, vol. 1, p. 399. Bonnet, 1955, Bibliographia Araneorum, vol.
2, p. 1255.
Types. Types of Theridion stictum from "roots of heath near
Winchester, and at Bloxworth, Dorset," England. Two male
syntypes of Crustulina borealis and four female syntypes of
Crustulina pallipes from Olympia, Washington, in the Museum
of Comparative Zoology.

Description. Carapace, sternum, brown; legs lighter brown.
Abdomen (Fig. 3) colored as in Steatoda borealis or lacking
a pattern ; sometimes white or dark brown. Epigynum illustrated
by Figure 2, male palpus by Figure 7. The openings of the
epigynum of a Wisconsin specimen are not connected by a fold.
Total length of females 2.2-2.7 mm. A female from Utah meas-
ured : total length, 2.6 mm. Carapace 1.06 mm. long, 1.04 mm.
wide. First femur, 1.14 mm. ; patella and tibia, 1.17 mm. ;
metatarsus 0.71 mm. ; tarsus, 0.57 mm. Second patella and tibia,
0.91 mm. ; third, 0.78 mm. ; fourth, 1.01 mm. Total length of
males 1.9-2.7 mm. A male from Utah measured: total length, 2.2
mm. Carapace 1.10 mm. long, 0.92 mm. wide. First femur,
1.04 mm.; patella and tibia, 1.10 mm.; metatarsus, 0.66 mm.;

372 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

tarsus 0.51 mm. Second patella and tibia, 0.88 mm. ; third, 0.66
mm. ; fourth, 0.93 mm.

Natural History. This species has been collected from leaf lit-
ter and from under boards and stones, often in moist places
(Kaston, 1948; Locket and Millidge, 1953). Roth has collected
it in fir needles in Oregon.

Distribution. England and parts of France (Simon, 1881;
Locket and Millidge, 1953), United States to Alaska (Map 1).
Judging by its wide distribution, this species is probably native
to North America.

Records. Alaska: (Chamberlin and I vie, 1947). Newfound-
land: (Hackman, 1954). Nova Scotia: Annapolis Co.; Kings
Co. Quebec: Gaspe Pen.; Brion Isl. ; Niapisca Isl. Ontario: 36
mi. N. of North Bay; Ottawa. Manitoba: Riding Mtn. British
Columbia: (Emerton, 1920) ; Departure Bay; Vanderhoof ; Well-
ington. Maine: Aroostook Co. ; Cumberland Co. ; Mt. Desert Isl. ;
New Hampshire: Carroll Co.; Mt. Washington. Massachusetts:
(Emerton, 1930) ; Essex Co.; Middlesex Co.; Plymouth Co. Con-
necticut: (Kaston, 1948). New York: (Crosby and Bishop,
1928); Rochester; Suffolk Co. New Jersey: Bergen Co. West
Virginia: Pendleton Co. Michigan. Illinois: (Kaston, 1955).
Minnesota: Aitkin Co. Nebraska: Lancaster Co. Texas: Hidalgo
Co. ; Matagorda Co. Wj/omm^; Yellowstone Natl. Pk. Colorado:
Fort Collins. Idaho: Oneida Co. Utah: Salt Lake City; Uintah
Co. Arizo7ia: Coconino Co. Washingto7i: (Worley, 1932) ;
Island Co. ; Olympia ; Pacific Co. ; San Juan Co. ; Seattle ; Sno-
homish Co.; Yakima. Oregon: Benton Co.; Coos Co.; Clackamas
Co.; Columbia Co.; Jackson Co.; Yamhill Co. California: Ala-
meda Co. ; Lassen Co. ; Los Angeles Co. ; Mendocino Co. ; Placer
Co.; San Francisco; Santa Barbara; Yolo Co. Baja California:
Islas Coronado ; ' ' San Jose. ' '

Ckustulina altera Gertsch and Archer
Figures 4-6, 8-10 ; Map 2.

Steatoda guttata, Emerton, 1882, Trans. Connecticut Acad. Sci., vol. 6, p.

20, pi. 4, fig. 2, $ . Emerton, 1902, The common spiders, p. 120, figs.

280-281,$. Not Cruslulina guttata (Wider).
Crustulma sticta, Keyserling, 1886, Die Spinnen Amerikas, vol. 2, pt. 2, p.

37, pi. 12, fig. 154, $,6. Not Crustulina sticta (O.P. Cambridge).

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA

373

Crustulina guttata, Banks, 1896, Jour. New York Ent. Soc, vol. 4, p. 191;
1900, Proc. Acad. Nat. Sci. Philadelphia, p. 533. Banks, 1910, Bull.
U.S. Natl. Mus. no. 72, p. 21 ; 1911, Proc. Acad. Nat. Sci. Philadelphia,
vol. 63, p. 445. Comstoek, 1912, The spider book, p. 360. Barrows,
1918, Ohio Jour. Sci., vol. 18, p. 303. Emerton, 1919, Ent. News, vol.
30, p. 167. Bishop and Crosby, 1926, Jour. Elisha Mitchell Sci. Soc,
vol. 41, p. 176. Crosby and Bishop, 1928, Mem. Cornell Agr. Exp. Sta.,

C. altera

Map 2. Distribution of Crustulina altera Gertsch and Archer.

no. 101, p. 1039. Emerton, 1930, Publ. Nantucket Maria Mitchell
Assoc, vol. 3, p. 164. Crosby and Bishop, 1936, Jour. New York Ent.
Soc, vol. 44, p. 44. Kaston, 1938, Bull. Connecticut Geol. Nat. Hist.
Surv., no. 60, p. 185. Comstoek, 1940, The spider book, rev. ed., p. 375.
Lowrie, 1948, Ecology, vol. 29, p. 336. Dowdy, 1950, Amer. Midland
Nat., vol. 43, p. 667. Bonnet, 1955, Bibliographia Araneorum, vol. 2,
p. 1256, in part. Not Crustulina guttata (Wider).
Crustulina altera Gertsch and Archer, 1942, Amer. Mus. Novitates, no. 1171,

374 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

p. 1, fig. 9,?. Chamberlin and Ivie, 1944, Bull. Univ. Utah, biol. ser.,
vol. 8, no. 5, p. 39, Muma, 1945, Bull. Univ. Maryland Agr. Exp. Sta.,
no. A38, p. 25. Archer, 1946, Paper Alabama Mus. Nat. Hist., no. 22,
p. 20. Kaston, 1948, Bull. Connecticut Geol. Nat. Hist. Surv., no. 70,
p. 74, figs. 30-33, 2119,9,^. Kaston, 1953, How to know the spiders,
p. 161, fig. 403,9. Levi and Field, 1954, Amer. Midland Nat., vol. 51.
p. 442. Kaston, 1955, Trans. Illinois Acad. Sci., vol. 47, p. 166.
Types. Male liolotype aud female allotype of C. altera from
Norwalk, Connecticut, in the American Museum of Natural His-
tory.

Description. Carapace, sternum orange-brown ; legs yellowish.
Abdomen (Fig. 10) with brown maculations sometimes similar
to that of C. sticta. Epigynum illustrated by Figure 5, male
palpus by Figure 8. Total length of females 1.5-2.0 mm. A
female from Illinois measured: total length 1.8 mm. Carapace
0.42 mm. long, 0.80 mm. wide. First patella and tibia 0.85 mm. ;
second, 0.67 mm. ; third, 0.49 mm. Fourth femur, 0.85 mm. ; patella
and tibia, 0.89 mm. ; metatarsus, 0.45 mm. ; tarsus, 0.41 mm. Total
length of males 2.0-2.3 mm. A male from Illinois measured : total
length 2.3 mm. ; carapace 1.23 mm. long, 0.98 mm. wide. First
femur 1.07 mm. ; patella and tibia, 1.18 mm. ; metatarsus, 0.72
mm. ; tarsus, 0.57 mm. Second patella and tibia, 0.98 mm. ; third,
0.75 mm.; fourth, 1.03 mm.

Natural History. This species is found under logs and stones.
In Wisconsin it is commonly collected by sifting leaf litter from
relatively dry forests. In Connecticut males overwinter as
adults. Egg sacs are 3 mm. in diameter made of few white
threads and containing about five pink eggs (Kaston, 1948).
Both Emerton (1919) and Crosby and Bishop (1939) collected
it with ballooning spiders.

Distribution. Eastern United States (Map 2). There are many
records from the "West and Canada ; however these seem to be
C. sticta.

Records. Massacliuseits: (Emerton, 1930) ; Boston. Connecti-
cut: (Kaston, 1948). New York: (Crosby and Bishop, 1928;
Gertsch, 1942). New Jersey: Cape May; Mercer Co. Maryland:
(Muma, 1945). District of Columbia: Washington. Virginia:
Fairfax Co. North Carolina: Durham; Buncombe Co.; Hay-
wood Co. South Carolina: Stumphouse Mtn. Georgia: (Cham-
berlin and Ivie, 1944). Florida: (Gertsch, 1942); Volusia Co.

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 375

Alabama: (Gertsch, 1942; Archer, 1946). Mississippi: Lauder-
dale Co. Louisiana: (Crosby and Bishop, 1936). Ohio: (Bar-
rows, 1918). Indiana: (Lowrie, 1948) Vanderburgh Co. Ten-
nessee: (Gertsch, 1942). Wisconsin: (Levi and Fiekl, 1954);
Crawford Co. ; La Crosse Co. ; Jackson Co. ; Sauk Co. ; "Waushara
Co. Illinois: Bond Co.; Jackson Co.; Macoupin Co.; Piatt Co.;
Union Co.; Will Co. Minnesota: Minneapolis. Missouri: (Dow-
dy, 1950) ; St. Louis. Arkansas: Yell Co.: Roaring River State
Pk.

Steatoda Sundevall

Steatoda Sundevall, 1833, Conspectus Arachnidum, p. 16. Type species:

" Theridion 4-punctatum" (= Steatoda bipunctata Linnaeus).
Asagena Simdevall, 1833, op. cit., p. 19. Type species: Asagena phalerata

Panzer.
Lithyphantes Thorell, 1869, On European spiders, p. 94. Type species:

Lithyphantes corollatus Linnaeus (= Steatoda albomaculata De Geer).
Teutana Simon, 1881, Les Araehnides de France, vol. 5, p. 161. Type

species: Teutana triangidosa Walckenaer.
Stearodea F.O.P. Cambridge, 1902, Ann. Mag. Nat. Hist., ser. 7, vol. 9,

p. 193. Type species: Aranea bipunctata Linnaeus.
Remarks on nomenclature. Sundevall listed four species as
belonging to Steatoda, but did not indicate the type. C.L. Koch
in 1845, according to F.O.P. Cambridge, 1902, withdrew Aranea
4-punctatum, the first species listed, from Steatoda and placed
it in Eucharia. Cambridge believed that one of the remaining
species, Theridion lunatum ought to be the type. If we were to
follow Cambridge, Achaea Keyserling (= Achaearanea Strand)
would be a subjective synonym of Steatoda. However, it seems
best to continue the usage of the last fifty years and assume
Aranea hipunctata to be the type.

Description. Medium sized to large theridiid spiders (2-11
mm. total length). Carapace slightly longer than wide in larger
species, longer than wide in smaller ones, sometimes with a
circular or transverse depression. Anterior eye row straight to
procurved when viewed from in front, posterior row straight
or slightly recurved when viewed from above. Size of eyes
variable. Carapace highest at posterior eye row. Chelicerae
about as long as height of carapace with one tooth on anterior
margin (Figs. 63, 87), or two {S. moesta). Legs of medium

376 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

length. First or fourth longest sometimes subequal in length,
third shortest. A distinct comb present on fourth tarsus. Ab-
domen subglobular in larger species, longer than wide or high
in smaller species. Colulus large (Figs. 62, 86).

Sexual dimorphism. Male slightly smaller than female. Cara-
pace and sternum of males sometimes more rugose ; carapace of
male with a stridulating organ on each side of pedicel. Anterior
end of abdomen above pedicel with a semicircular sclerotized
carina which apparently rubs against stridulating organ. Males
of many species with chelicerae enlarged, and endites swollen,
sometimes with tubercles on endites. The males of some species
(S. americana) may have the first pair of legs thicker, and the
second with a spine. Males are much less common in collections
than females.

Genitalia. Epigynum often with a flat plate and a median lobe
projecting posteriorly. Two seminal receptacles present. Ducts
heavily sclerotized in the 8. Mpunctata group. Palpus with a
large radix (R in Figs. 60, 61, 69, 75, 153, 154), median apoph-
ysis (M), conductor (C) and embolus (E). Embolus may
have an elaborate base which may be of diagnostic value in both
the S. fulva and 8. hipunctata group. Paracymbial hook (P)
on the raesal side of alveolus in some species (Fig. 60) in others
distal or ectal apparently holding median apophysis in all
species. In 8. alhomaculata which has the paracymbial hook
ectal, the median apophysis has a process which probably fits
against this structure (Fig. 61).

Diagnosis. This genus differs from other theridiid genera by
having a large colulus ; one or two teeth on the anterior margin
of the chelicerae, no teeth on the posterior. It differs from
Latrodectus in that the lateral eyes are touching each other or
are separated by less than their diameter ; from Crustulina in
lacking a spur on the mesal side of the cymbium of male palpus
and lacking a sclerotized ring on the abdomen of females around
the pedicel.

Distribution. Members of this genus are found in all parts
of the world. Several are cosmopolitan.

Misplaced species. The following North American species de-
scribed as Lithyphantes or Steatoda are misplaced.

Lithyphantes cyaneus Worley, 1928, Ann. Ent. Soc. Amer.,
vol. 21, p. 621, figs. 5,6,2, is a 8inga (Argiopidae).

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 377

Lfithyphantes mimoides Chamberlin, 1919, Jour. Ent. Zoo.,
vol, 12, p. 8, pi. 3, fig. 4, 9 , is Enoplognatha marmorata
(Hentz).

Lithyphantes oophorus Petrunkevitch, 1930, Trans. Connecti-
cut Acad. Sci., vol. 30, p. 170, figs. 8, 9, 9 , is Coleosoma floridana
Banks.

Steatoda fusca Emerton, 1894, Trans. Connecticut Acad. Sci.,
vol. 9, p. 407, pi. 2, fig. 1, 9 , (5 is a Ctenium.

Steatoda nigra Emerton, 1884, Trans. Connecticut Acad. Sci.,
vol. 6, p. 21, pi. 4, fig. 4:,9,$ is a Dipoena.

F.O.P. Cambridge (1902, Biologia Centrali Americana, Ara-
neidea, vol. 2) described several species of Theridion in the genus
Steatoda.

Steatoda niqrofemorata (Keyserling), new combination

Figures 11-13; Map 3.

Lithyphantes nigrofemoratus Keyserling, 1882, Die Spinnen Amerikas, vol.
2, no. 1, p. 139, pi. 6, lig. 87,9. F.O.P. Cambridge, 1902, Biologia
Centrali Americana, Araneidea, vol. 2, p. 376, pi. 35, fig. 14,9. Pet-
runkevitch, 1911, Bull. Amer. Mus. Nat. Hist., vol. 29, p. 183. Chamber-
lin, 1916, Bull. Mus. Comp. Zool., vol. 60, p. 232. Banks, 1929, ibid.,
vol. 69, p. 86. Mello-Leitao, 1941, Kev. Mus. La Plata, vol. 2, p. 141.
Roewer, 1942, Katalog der Araneae, vol. 1, p. 408.

Asagen<i alticeps Kej-serling, 1886, Die Spinnen Amerikas, vol. 2, pt. 2, p.
4, fig. 136, $ . Simon, 1894, Histoire naturelle des Araignees, vol. 1,
p. 576. F.O.P. Cambridge, 1902, Biologia Centrali Americana.
Araneidea, vol. 2, p. 377, pi. 35, fig. 16, cJ. Simon, 1903, Histoire
naturelle des Araignees, vol. 2, p. 990. Petrunkevitch, 1911, Bull. Amer.
Mus. Nat. Hist., vol. 29, p. 170. Eeimoser, 1939, Ann. Naturhist. Mus.
Wien, vol. 50, p. 344. Mello-Leitao, 1940, Arq. Zool., vol. 12, p. 176.
Roewer, 1942, Katalog der Araneae, vol. 1, p. 396. Bonnet, 1955,
Bibliographia Araneorum, vol. 2, p. 752. Caporiacco, 1955, Acta Biol.
Venezuelica, vol. 1, p. 329, fig. 22,9. NEW SYNONYMY.

lAthxjphantes lattus O.P. Cambridge. 1896, Biologia Centrali Americana,
Araneidea, vol. 1, p. 181, pi. 22, fig. \2,$. NEW SYNONYMY.

lAsagena patagonica Tullgren, 1901, Svenska Exped. Magellanslanderna,

vol. 2, no. 10, p. 194, pi. 15, fig. 4,9- Petrunkevitch, 1911, Bull. Amer.

Mus. Nat. Hist., vol. 29, p. 171. Roewer, 1942, Katalog der Araneae,

vol. 1, p. 397. NEW SYNONYMY.

Types. Female holotype of Lithyphantes nigrofemoratus from

Monte Rico. Peru, in the collection of the University of War-

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saw, Poland. Male hole type of Asageria alticeps from Nueva
Granada, Colombia. Male liolotype of Lithyphantes laetus from
Costa Rica in the British Museum (Natural History). Three
female syntypes of Asagena patagonica from Patagonia, prob-
ably in the Riksmuseum, Stockholm.

Description. Carapace, sternum, legs yellow-brown; femora
much darker, ends of tibiae darker. Abdomen black, dorsum
usually with two transverse lines, an anterior one and one
posterior which is broken on each side. South American speci-
mens often with spots on dorsum ; venter with a white spot
behind genital groove. Anterior median eyes smaller than

S. nigrofemorofa

Map 3. Distribution of Steatoda nigrofemoraia (Keyserling).

others, one diameter apart, one-half to one diameter from
laterals. Posterior median eyes three-quarters their diameter
apart, a little more than one diameter from laterals. Male cheli-
cerae enlarged, lacking distinct tooth on anterior margin. Epig-
ynum with a wrinkled scape (Fig. 13), male palpus illustrated
by Figure 11. The embolus of the palpus is sometimes broken
off. Total length of females 3.8-4.3 mm. A female from Panama
measured: total length, 4.2 mm. Carapace 1.6 mm. long, 1.2 mm.

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 379

wide. First patella and tibia, 1.7 mm.; second, 1.5 mm.; third,
1.3 mm. Fourth femur, 1.5 mm. ; patella and tibia, 1.8 mm. ;
metatarsus, 1.1 mm. ; tarsus, 0.7 mm. Total length of males
3.3-3.5 mm. A male from Panama measured total length, 3.5
mm. Carapace 1.8 mm. long, 1.1 mm. wide. First patella and
tibia, 1.6 mm.; second, 1.5 mm.; third, 1.2 mm. Fourth femur,
1.6 mm. ; patella and tibia, 1.9 mm. ; metatarsus, 1.3 mm. ;
tarsus, 0.7 mm. Tullgren's specimens were larger, total length
6 mm.

Natural History. This species has been collected under a
fallen tree in Peru. Tullgren found his in dry, sandy localities.

Distribution. South America, north to Veracruz (Map 3).

Records. Veracruz: 10 mi. S. of San Jose del Carmen. Guate-
mala: Quirigua. Nicaragua: Musawas, AYaspuc River. Costa
Rica: (Reimoser, 1939). Panama: Barro Colorado Island; Bo-
quete; La Compana; Santa Ro.sa, Colon Prov. Venezuela:
(Caporiacco, 1955). British Guiana: (Mello-Leitao, 1940). Peru:
(Chamberlin, 1916); Ayacucho ; Huanuco; Huancayo; Tingo
Maria; Pucara. Brazil: Minas Geraes; Minha Serinha Diaman-
tina. Parana: Cavinna; Rolandia. Argentina: (Tullgren, 1901;
Mello-Leitao, 1941) ; Salta.

Steatoda moesta (O.P. Cambridge), new combination
Figures 15-18 ; Map 4.

Steatoda brasiliana, Keyserling, 1886, Die Spinnen Ameriias, vol. 2, pt. 2,
p. 238, pi. 20, fig. 293,$. Not Steatoda brasiliana Keyserling, 1884,
ibid., pt. 1, p. 115, pi. 5, fig. 75,^.

Asagena moesta O.P. Cambridge, 1896, Biologia Centrali Americana.
Araneidea, vol. 1, p. 209, pi. 25, fig. 4,$. F.O.P. Cambridge, 1902, ibid.,
vol. 2, p. 35, fig. 18,5. Petrunkevitch, 1911, Bull. Amer. Mus. Nat.
Hist., vol. 29, p. 170. Eoewer, 1942, Katalog der Araneae, vol. 1, p.
397. Bonnet, 1955, Bibliographia Araneorum, vol. 2, p, 753. Kraus,
1955, Abhandl. Seuckenbergischen Naturf. GeselL, no. 493, p. 15.

Type. Female type of Asagena moesta from Guatemala in
the British Museum (Natural History).

Description. Carapace, sternum dark brown. Legs dark
brown, except fourth, which have femora and tibiae orange
brown, with ends darker. Abdomen black with white marks on
dorsum (Fig. 15) ; venter with white marks on each side of

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epigastric area. Male with sclerotized portions orange brown.
Eyes subequal in size. Anterior median eyes separated by two-
thirds their diameter. Posterior medians two-thirds diameter
apart, two-thirds to one diameter from laterals. Chelicerae with
two teeth on anterior margin. Epigynum a black spot in a lighter
area, posterior to which is a small scape (Fig. 17). In South
American specimens the black spot may be larger, the posterior
scape indistinct. Coloration and shape quite variable. Male
palpus illustrated by Figure 18. Total length of females 4.8-8.7
mm. A female from Panama measured total length, 8.7 mm.
Carapace 3.8 mm. long; 3.3 mm. wide. First femur, 5.0 mm.;

Map 4. Distribution of Steatoda mo.esta (O.P. Cambridge) and S. salten-
sis, new species.

patella and tibia, 5.4 mm. ; metatarsus, 4.0 mm. ; tarsus, 1.8 mm.
Second patella and tibia, 4.3 mm. ; third, 3.5 mm. ; fourth, 4.9 mm.
Total length of males, 6.2-8.0 mm. A male from Panama meas-
ured total length, 6.5 mm. Carapace 3.4 mm. long; 2.5 mm.
wide. First femur, 4.3 mm. ; patella and tibia, 4.3 mm. ; meta-
tarsus, 3.3 mm. ; tarsus, 1.6 mm. Second patella and tibia, 3.6
mm. ; third, 2.7 mm. ; fourth, 4.2 mm.

Distribution. From Tabasco to Brazil, Map 4.

Becords. Ta&a.sco: Teapa (C. and M. Goodnight), $. Chiapas:
Rio Huixtla, N. of Huixtla (T. C. Schneirla), 9 . El Salvador:
(Kraus, 1955). Panama: El Valle (A. M. Chickering), 9.^ ;

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 381

Barro Colorado Island (A. M. Chickering), 9 , 5 . Peru: Dept.
Huanueo; Acomayo, 2160 in. (F. AVoytkowski), 9 . Brazil:
Est. Rio de Janeiro: Teresopolis (H. Sick), 9 .

Steatoda quaesita (O.P. Cambridge), new combination

Figure 14.

Liihyphantes quaesitus O.P. Cambridge, 1896, Biologia Central! Americana,
Araneidea, vol. 1, p. 180, pi. 22, fig. 6,S. F.O.P. Cambridge, 1902,
ibid., vol. 2, p. 376, pi. 35, fig. 13, $. Petrunkevitch, 1911, Bull. Amer.
Mus. Nat. Hist., vol. 29, p. 183. Roewer, 1942, Katalog der Araneae,
vol. 1, p. 408.
Type. Male holotype from Teapa, Tabasco, in the British
Museum (Natural History).

Only the palp of the type was examined. It is probably the
male of either *§. salfensis or more likely that of S. autumnalis.

, Steatoda saltensis, new species
Figures 19, 20 ; Map 4.

Lithyphantes lugubris F.O.P. Cambridge, 1902, Biologia Centrali Americana,
Araneidea, vol. 2, p. 376 (in part), pi. 35, fig. 15c (not fig. 15b).

Types. Female holotype and one female paratype from forty
miles northeast of El Salto, Durango, Mexico, Aug. 11, 1947
(AY. J. Gertsch), in the American Museum of Natural History.

Description. Carapace, sternum, legs brown. Abdomen black
with a light transverse anterior band ; venter with a white line
behind genital furrow. Eyes subequal in size. Anterior median
eyes one diameter apart, three-quarters of a diameter from
laterals. Posterior median eyes one diameter apart, one and
three-quarters diameters from laterals. Laterals separated by
three-quarters diameter. Total length of females 7.6-10.3 mm.
The holotype measured total length 10.3 mm. Carapace 3.7
mm. long ; 3.0 mm. wide. First femur, 4.3 mm. ; patella and
tibia, 4.8 mm. ; metatarsus, 3.7 mm. ; tarsus, 1.4 mm. Second
patella and tibia, 4.0 mm. ; third, 3.0 mm. ; fourth, 4.7 mm.

Diagnosis. Only details of the epigynum (Fig. 20) distinguish
this species from the related Steatoda autumnalis.

Distribution. Central Mexico (Map 4). A female paratype of
L. luguhris which F.O.P. Cambridge thought to be immature

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was collected in Bugaba, Panama. Recent Panamanian collectors
have not included S. saltensis again.

Records. Durango: 30 mi. NE. of El Salto, Aug. 11, 1947
(W. J. Gertsch), 9 . Distrito Federal: Contreras, May 30, 1946,
8000-9000 ft. (J.C. and D.L. Pallister), 9 . 'IZacatecas: Tena-
yuca, July 14, 1943 (C. Bolivar), 9.

Steatoda autumnalis (Banks), new combination
Figures 21, 22 ; Map 5.

Lithyphantcs autumnalis Banks, 1898, Proc. California Acad. Sci., 3rd. ser.,
vol. 1, p. 240, pi. 14, fig. 6. r.O.P. Cambridge, 1902, Biologia Centrali
Americana, Araneidea, vol. 2, p. 376. Petrunkevitch, 1911, Bull. Amer.
Mus. Nat. Hist., vol. 29, p. 182. Eoewer, 1942, Katalog der Araneae
vol. 1, p. 407.

LUhyphant.es clams O.P. Cambridge, 1898, Biologia Centrali Americana,
Araneidea, vol. 1, p. 252, pi. 38, fig. 6. F.O.P. Cambridge, 1902, ihid.,
vol. 2, p. 376. Petrunkevitch, 1911, Bull. Amer. Mus. Nat. Hist., vol.
29, p. 182. Roewer, 1942, Katalog der Araneae, vol. 1, p. 407. NEW
SYNONYMY.

Map 5. Distribution of Steatoda autumnalis (Banks).

Lithyphantes lugubris F.O.P. Cambridge, 1902, Biologia Centrali Americana,
Araneidea, vol. 2, p. 376, (in part), pi. 35, fig. 15b (not 15c). Petrunke-
vitch, 1911, Bull. Amer. Mus. Nat. Hist., vol. 29, p. 183. 1 Banks,
1913, Proc. Acad. Nat. Sci. Philadelphia, p. 678. ? Reimoser, 1937,
Ann. Naturhist. Mus. Wien, vol. 50, p. 344. Roewer, 1942. Katalog
der Araneae, vol. 1, p. 408. NEW SYNONYMY.
Types. Immature, less than half grown, holotype of Lithy-
phantes autumnalis Banks, from Tepic, Nayarit, in the Museum

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 383

of Comparative Zoology. Immature holotype of Lithyphantes
clarus from Patzcuara, Michoacan. Female holotype of Lithy-
phantes lugubris from Omiltemi, Guerrero. The last two are
in the British Museum (Natural History).

Remarks on nomenclature. Although the types of Lithy-
phantes autumnalis and L. clarus can not be determined with
absolute certainty, L. autumnalis has the typical pattern of the
juvenile of this species, and adults have been collected in the
type locality. L. clarus is probably a penultimate female, judg-
ing by its coloration. The type locality is well within the range
of this species. The publication date of L. autumnalis is May
28, 1898 ; that of L. clarus, June, 1898.

Description. Carapace, sternum, legs dark brown. Abdomen
black with a white stripe around anterior margin of dorsum;
venter with a short white transverse line behind genital groove.
Immatures seem to have abdomen brightly colored with patches
of light and dark pigment. Eyes on slight tubercles ; anterior
medians slightly smaller than others, one diameter apart, and
one diameter from laterals. Posterior median eyes one diameter
apart, one and one-half diameters from laterals. Epigynum
illustrated by Figure 22. Total length of females 7.8-11.0 mm.
A female measured : total length 8.7 mm. Carapace 3.7 mm.
long, 3.0 mm. wide. First femur, 4.0 mm. ; patella and tibia,
4.9 mm. ; metatarsus, 3.5 mm. ; tarsus, 1.3 mm. Second patella
and tibia, 3.3 mm. ; third, 3.0 mm. ; fourth, 4.4 mm. The male
is not known.

Distril)utw7i. Central Mexico. Map 5. The records of Costa
Rica and Panama (F.O.P. Cambridge, 1902; Banks, 1913; Rei-
moser, 1939) are probably erroneous.

Records. Nayarit: Tepic. Jalisco: Nevado de Colima; Tlaque-
pague; W. side of Lake Sayula; Tizapan. Mexico: Ixtapan de la
Sal. Distrito Federal: Santa Rosa. Morelos: Oaxtepec; Acat-
lipa; Cuantla. Tlaxcala: Tlaxcala. Puehla: Tehuacan. Guer-
rero: Taxco; 23 mi. S. of Chilpancingo.

Steatoda transversa (Banks), new combination

Figures 23-27 ; Map 6.

Lithyphantes transversus Banks, 1898, Proc. California Acad. Sci., 3rd ser.,
vol. 1, p. 240, pi. 14, fig. 5,9. Petrunkevitch, 1911, Bull. Amer. Mus.

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BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

Nat. Hist., vol. 29, p. 183. Roewor, 1942, Katalog der Araneae, vol. 1,
p. 408.

Type. One female from El Taste, Baja California, in the
Museum of Comparative Zoology.

Description. Carapace, sternum, legs, orange-brown. Abdomen
dark purplish with white spots on dorsum and indistinct white
spots on venter. Anterior lateral eyes largest, posterior laterals
slightly smaller, median eyes subequal and smallest. Anterior
medians three-quarters diameter apart, the same distance from
laterals. Posterior medians half their diameter apart, one di-
ameter from laterals. Chelicerae of male slightly swollen and
endites enlarged. Epigynum (Pig. 24) with two dark spots

S. tronsverso

Map 6. Distribution of Steatoda transversa (Banks).

slightly more than their diameter apart. Male palpus illustrated
by Figures 26, 27. A female from Arizona measured : total
length, 4.3 mm. Carapace 1.6 mm. long, 1.3 mm. wide. First pa-
tella and tibia, 1.4 mm. ; second, 1.2 mm.; third, 1.0 mm. Fourth
femur, 1.3 mm. ; patella and tibia, 1.7 mm. ; metatarsus, 0.9 mm. ;
tarsus, 0.7 mm. A male from Arizona measured : total length,
3.1 mm. Carapace 1.6 mm. long, 1.1 mm. wide. First patella
and tibia, 1.3 mm.; second, 1.1 mm.; third, 0.9 mm. Fourth
femur, 1.2 mm. ; patella and tibia, 1.4 mm. ; metatarsus, 0.9 mm. :
tarsus, 0.6 mm.

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 385

Natural History. Roth collected this species in Arizona by
sweeping an alfalfa field.

Distribution. Texas, Arizona to Oaxaca. Map 6.

Records. Texas: Hays Co.; Hidalgo Co.: Edinburg. Arizona:
Yuma Co.: Yuma Mesa. California: Riverside Co.: Indio. So-
nera: 10 mi. S. of Hermosillo. Nayarit: Acaponeta; Jesus Maria.
Jalisco: Plan de Barrancas. Colima: 20 mi. N. ; 7 mi. S. of
Colima; Manzanillo. Morelos: Alpuyecca. Veracruz: Plan del
Rio. Oaxaca: Oaxaca.

Steatoda quadrimaculata (O.P. Cambridge),

new combination

Figures 28-31; Map 7.

Asag.ena quadrimaculata O.P. Cambridge, 1896, Biologia Central! Ameri-
cana, Araneidea, vol. 1, p. 189, pi. 23, fig. 12,5. Simon, 1897, Proc.
Zool. Soc. London, p. 864. F.O.P. Cambridge, 1902, Biologia Central]
Americana, Araneidea, vol. 2, p. 378, pi. 35, fig. 17,3. Petrunkevitch,
1911, Bull. Amer. Mus. Nat. Hist., vol. 29, p. 171. Chiekering, 1936,
Trans. Amer. Micros. Soc, vol. 55, p. 451. Bryant, 1940, Bull. Mus.
Comp. Zool., vol. 86, p. 302. Roewer, 1942, Katalog der Araneae, vol.
1, p. 397. Bonnet, 1955, Bibliographia Araneorum, vol. 2, p. 757.
Lithyphantes hermosa Banks, 1909, Proc. Acad. Nat. Sci. Philadelphia, vol.
61, p. 205, pi. 6, fig. 44,9. Petrunkevitch, 1911, Bull. Amer. Mus. Nat.
Hist., vol. 29, p. 182. Reimoser, 1939, Ann. Naturhist. Mus. Wien,
vol. 50, p. 344. Roewer, 1942, Katalog der Araneae, vol. 1, p. 407.
NEW SYNONYMY.
Lithyphantes tricolor Chamberlin and Ivie, 1944, Bull. Univ. Utah, biol. ser.,

vol. 8, no. 5, p. 44, figs. 67-74, $ . NEW SYNONYMY.
Asagena marcuzzii Caporiacco, 1955, Acta Biol. Veuezuelica, vol. 1, p. 330,
fig. 23,9. NEW SYNONYMY.
Types. Male holotype of Asagena quadrimaculata from An-
tigua, Guatemala, in the British Museum (Natural History).
Female holotype of Lithyphantes hermosa from Punta Arenas,
Costa Rica, in the Museum of Comparative Zoology. Female
holotype of Lithyphantes tricolor from Savannah Beach, Geor-
gia, in the American Museum of Natural History. Female holo-
type of Asagena marcuzzii from Maiquetia, D.F., Venezuela, in
the Museo de Biologia, Universidad Central di Venezuela.
Description. Carapace, sternum, legs yellow to orange, some

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duskiness on femora. Abdomen dark purple with 6 or 7 dorsal
white spots, a U-shaped light spot on venter and smaller spots
on each side. Anterior median eyes slightly smaller than others,
separated by three-quarters diameter, one-third diameter from
laterals. Posterior medians separated by one-third diameter, one-
half to one diameter from laterals. Lateral eyes not separated.
Chelieerae and endites of males enlarged. Epigynum with a
dark spot each in a pair of depressions (Fig. 29). Male palpus
illustrated by Figures 30, 31. Total length of females 2.7-4.0
mm. A female from Tamaulipas measured total length, 3.6 mm.
Carapace 1.36 mm. long, 0.98 mm. wide. First patella and tibia,
1.35 mm.; second, 1.13 mm.; third, 0.95 mm. Fourth femur,
1.17 mm., patella and tibia, 1.42 mm. ; metatarsus, 0.78 ; tarsus,
0.52 mm. Total length of males, 2.2-3.2 mm. A male from

G5?-"I

S quodrimaculato

Map 7. Distribution of Steatoda quadrimaculata (O.P. Cambridge).

Tamaulipas measured : total length, 3.1 mm. Carapace 1.60 mm.
long, 1.14 mm. wide. First patella and tibia, 1.54 mm.; second,
1.36 mm.; third, 1.10 mm. Fourth femur, 1.40 mm.; patella
and tibia, 1.74 mm.; metatarsus, 1.20 mm.; tarsus, 0.60 mm.

Distribution. Southern United States to northern South
America. Map 7.

Records. Florida: Alachua Co.; Collier Co.; Highlands Co.
Alabama: Baldwin Co. Mississippi: Covington Co. Louisiana:
Beauregard Par. ; St. Helena Par. ; Shreveport. Texas: Chambers

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 387

Co., Hidalgo Co.; Starr Co. Tamaulipas: 20 mi. N. of Lemon;
2 mi. E. of Nuevo Morelos; 13-15 mi. S., 27 mi. N., 35 mi. N. of
Villa Juarez. San Luis Potosi: Tamazunchale ; 2 mi. N. of
Tamazunehale ; 5 mi. N., 10 mi. N., 13 mi. N. of Valles ; Valles.
Hidalgo: Chapulhuaean. Veracruz: Cordoba. Nayarit: Campo-
stela; La Libertad. Colima: 25 mi. E. of Mazamitla. Guerrero:
Lo Bajo; Arcelia; 32 mi. N. of Acapulco. Oaxaca: Palomares;
Soledad ; Tolosa; San Felipe; Tehiiantepec. Chiapas: Las
Cruzes. Is. Revillagigedo: Socorro Is. Panama: (Chickering,
1936). Cuba: (Bryant, 1940). Lesser Antilles: (Simon, 1897).
Venezuela: (Simon, 1897).

Steatoda fulva group

This group is represented by three similar species which are
sympatrie over most of their ranges.

Description. Carapace, sternum, legs orange to brown ; some-
times legs red-brown with distal segments lighter. Abdomen
purplish brown with dorsal white spots (Figs. 51, 52, 54, 55),
venter with a white spot posterior to the genital groove, another
anterior to spinnerets, both frequently connected by a white line.
8. pulcher may be differently colored. In all three species eyes
subequal in size ; or frequently anterior medians larger than
others, rarely slightly smaller. Anterior medians one-half to one
diameter apart, one-quarter to one-half their diameter from
laterals. Posterior eyes equi-distant, two-thirds to one diameter
apart. Lateral eyes slightly separated or touching each other.
The genitalia and measurements are similar.

Variation. The coloration is variable in 8. medialis and 8.
pulcher. The epigyna of all three species, the palp of 8. medialis
and 8. pulcher are variable.

Natural History. Virtually nothing is known of the natural
history of these three species. It is assumed that all three are
found in arid situations under stones, logs and in debris.

Key to females of the Steatoda fulva group

1. Anterior edge of "transverse bridge" of epigynum pointing anterior;
"bridge" semicircular in cross-section (Figs. 43, 44) medialis

388 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

1. Anterior edge of "bridge" of epigynum, straight or pointing posterior.

(Figs. 46, 49) 2

2. Posterior edge of ' ' bridge ' ' with two lobes each as wide as inter-

mediate area, bridge flat in cross-section. (Fig. 49, 50) . . .pulcher
2. Posterior edge of ' ' bridge ' ' more or less straight with lobes widely

separated. (Figs. 46, 47) fulva

Key to males of the Steatoda fulva group

1. Palpus in ventral view with large portion of median apophysis showing
on mesal side, outside embolic tube (Figs. 34-36) medialis

1. Palpus in ventral view with only edge of median apophysis showing on

mesal side outside of embolic tube (Figs. 32, 38, 41) 2

2. Palpus in ventral view with area surrounded by embolic tube as high

as wide, subcircular ; basal membrane of embolus covering less than

one half of area enclosed by embolus (Figs. 32, 33) fulva

2. Palpus in ventral view with area surrounded by embolic tube elliptical,
much wider than high (Figs. 39, 41) ; or with area surrounded by
embolic tube subcircular with basal membrane of embolus covering
more than half of enclosed area (Fig. 38) pulcher

Steatoda medialis (Banks), new combination
Figures 34-36, 42-44, 53-55 ; Map 8.

Steatoda distincta Thorell, 1877, Bull. U.S. Geol. Surv., vol. 3, p. 485.

Banks, 1893, Jour. New York Ent. Soc, vol. 1, p. 124; ? 1895, Ann.

New York Acad. Sci., vol. 8, p. 423 ; 1910, Bull. U.S. Natl. Mus., no. 72,

p. 21. Petrunkevitch, 1911, Bull. Amer. Mus. Nat. Hist., vol. 29, p.

18S. '^ot Steatoda distincta (Blackwall).
Litltyphantes punctulata Marx, 1898, in Banks, Proc. California Acad. Sci.,

3rd ser., vol. 1, p. 239, pi. 14, fig. 4,$. Petrunkevitch, 1911, Bull.

Amer. Mus. Nat. Hist., vol. 29, p. 183. Roewer, 1942, Katalog der

Araneae, vol. 1, p. 408. NEW SYNONYMY.
Litltii pliant es medialis Banks, 1898, Proc. California Acad. Sci., 3rd. ser., vol.

1, p. 240, pi. 14, fig. 3,9. Banks, 1902, Proc. U.S. Natl. Mus., vol. 25,
p. 214. Banks, 1910, Bull. U.S. Natl. Mus., no. 72, p. 21. Petrunke-
vitch, 1911, Bull. Amer. Mus. Nat. Hist., vol. 29, p. 183.

Litlujphantes distincta, Chamlierlin, 1933, Bull. Univ. Utah, biol. ser., vol.

2, no. 2, p. 8, pi. 1, fig. 9,5. Gertsch, 1935, Amer. Mus. Novitates, no.
792, p. 21. Roewer, 1942, Katalog der Araneae, vol. 1, p. 408.

Types. Female type of Steatoda distincta from Colorado in
the Kiksmuseum, Stockholm. One female syntype and fragments
of two juveniles of Lithyphantes punctulata are alleged to have
come from Las Palmas, Baja California, in the Museum of Com-
parative Zoology. Two female syntypes of Lithyphantes medialis

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA

389

from San Jose del Cabo, Baja California, in the Museum of
Comparative Zoology.

Remarks^ on nomenclature. The name Steatoda distincta
Thorell is not available since Latrodectus distincta Blackwall,
1859 (Ann. Mag. Nat. Hist., ser. 3, vol. 4, p. 260) probably be-
longs to this genus and had been placed by Roewer (1942) in
Lithyphanies. Although Lithyphantes punctnlata has page
priority over L. medialis, Banks' name medialis is here used. I
doubt that the type specimens of L. punctidata came from Baja
California as indicated; they appear to come from the eastern

mediolis

Map 8. Distribution o/ Steatoda medialis (Banks). Squares: manr
spotted coloration; circles : few spotted coloration.

portion of its range. As the type locality is uncertain, the name
L. medialis seems preferable.

Description. Epigynum illustrated by Figures 43, 44; male
palpus by Figures 34-36. Total length of females 3.2-6.5 mm.
A female from California measured : total length 5.6 mm. Cara-
pace 2.1 mm. long, 1.7 mm. wide. First femur, 2.7 mm. ; patella
and tibia, 2.9 mm. ; metatarsiLS, 2.3 mm. ; tarsus, 0.9 mm. Second

390 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

patella and tibia, 2.3 mm. ; third, 1.9 mm. ; fourth, 2.9 mm. Total
length of males 2.9-5.8 mm. A male from California measured :
total length 5.4 mm. Carapace 2.4 mm. long, 1.6 mm. wide. First
femur, 3.0, mm. ; patella and tibia, 3.4 mm. ; metatarsus, 2.7 mm. ;
tarsus, 1.2 mm.; Second patella and tibia, 2.9 mm.; third, 2.4
mm. ; fourth, 3.4 mm.

Variation. There is considerable individual variation in the
shape of the epigynum, and relative eye size. The palpal tibia is
long in western specimens (Fig. 36), short in eastern ones (Fig.
34).

The difiference in dorsal marking is striking. Eastern speci-
mens commonly have four rows of spots on the dorsum of abdo-
men (Fig. 55), western ones have only one row of large spots
with one spot on each side (Fig. 54). Specimens from the center
of the range from Utah to central Mexico often lack spots. A
number of specimens from the Gulf of California had three lines
of spots. The type of L. punctulata is of the variety with four
rows of spots, L. medialis of the variety with one row. Map 8
shows the variety having four rows of spots as squares, the
fewer-spotted, or those lacking spots, by circles. Collections
having both patterns or the few individuals of intermediate
pattern are also indicated by circles.

Natural History. This species has been collected in Colorado
under stones in an arid area at 7000 feet elevation.

Distribution. Wyoming, Utah to Hidalgo. Map 8.

Records. Texas: Austin; El Paso ; Hidalgo Co.; Hudspeth
Co. ; Jeff Davis Co. ; Kerr Co. ; Llano Co. ; Presidio Co. ; Starr Co. ;
Terrell Co.; Webb Co.; Wise Co.; Zapata Co. Wyoming: Hawk
Springs, Goshen Co. Aug. 5, 1952 (B. Malkin). Colorado:
Chaffee Co. New Mexico: Eddy Co.; Luna Co.; San Miguel Co.
Utah: (Chamberlin and Ivie, 1933) ; Emery Co.; Salt Lake City;
Sevier Co.; Uintah Co.; AVayne Co. Arizona: (Banks, 1902;
Gertsch, 1935); Cochise Co.; Coconino Co.; Gila Co.; Organ
Pipe Natl. Mon. ; Phoenix ; Pima Co. ; Santa Cruz Co. ; Tucson.
Oregon: Malheur Co.: Malheur Riv. Canyon, Sept. 10, 1949 (V.
Roth). California: Berkeley; Contra Costa Co.; Inyo Co.; Los
Angeles Co. ; Mono Co. ; Riverside Co. ; San Bernandino Co. ; San
Diego Co.; San Nicolas Isl. ; Santa Barbara. Tamaulipas: 20
mi. E. of Villa Juarez; Arroyo Chorreras. Nuevo Leon: nr.
Monterey; 30 mi. E. of General Bravo; China. Coahuila: 5 mi.

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 391

W. of Saltillo; La Gloria. Chihuahua: Primavera, 5500-6000 ft.;
Huejotitlan ; 50 mi. S. Ahumacla ; 5 mi. S. Chihuahua ; 6 mi. S.
Gallego; La Sauceda, 7000 ft.; Valle de Olivos, 5000 ft. Sonora:
Punta Penasco ; 15 mi. W. Agiabambo ; Guaymas ; Navajoa ; Isla
Pelicano. Baja California: San Jose del Cabo; 45 mi. E. of
Tecate, 3600 ft. ; Isla. Cedros ; Isla. Habana ; Sal Si Puedes Isl. ;
Carman Isl.; Coronados Isl.; Smith's Isl.; La Paz; Santa Cruz
Isl. Durango: San Juan del Rio; La Loma. Sinaloa: 6 mi. S.
Caliacan. Nayarit. Jalisco: 20 mi. N. of La Quemada. Distrito
Federal: Teotihuacan. Hidalgo: Ixmiquilpan.

Steatoda fulva (Keyserling), new combination
Figures 32, 33, 45-47, 52 ; Map 9.

Lithyphatites fulvus Keyserling, 1882, Die Spinnen Amerikas, vol. 2, pt. 1,
pi. 6, fig. 89,$. Marx, 1889, Proc. U.S. Natl. Mus. vol. 12, p. 522.
Banks, 1899, Proc. Ent. Soc. Washington, vol. 4, p. 189; 1900, Proc.
Acad. Nat. Sci. Philadelphia vol. 52, p. 533; 1904, ihid., vol. 56, p. 653;
1910, Bull. U.S. Natl. Mus., no. 72, p. 21. Petrunkevitch, 1911, Bull.
Amer. Mus. Nat. Hist., vol. 29, p. 182. Comstock, 1912, The spider
book, p. 362, fig. 368,9. Worley and Pickwell, 1931, Univ. Nebraska
Studies, vol. 27, p. 27. Fox, 1940, Proc. Biol. Soc. Washington, vol.
53, p. 41. Comstock, 1940, The spider book, rev. ed., p. 377, fig. 368,$.
Chamberlin and Ivie, 1941, Bull. Univ. Utah, biol. ser., vol. 6, no. 3,
p. 11. Koewer, 1942, Katalog der Araneae, vol. 1, p. 408. Chamberlin
and Ivie, 1944, Bull. Univ. Utah, biol. ser., vol. 8, no. 5, p. 44.

Lithyphantcs parvula Banks, 1898, Proc. California Acad. Sci., 3rd. ser.,
vol. 1, p. 238, pi. 14, fig. 1,9,$. Petrunkevitch, 1911, Bull. Amer.
Mus. Nat. Hist., vol. 29, p. 183. Roewer, 1942, Katalog der Araneae,
vol. 1, p. 408. NEW SYNONYMY. Not Steatoda parvula Saito, 1933.

Lithyphantes p-ulcher, Banks, 1898, Proc. California Acad. Sci., 3rd. ser.,
vol. 1, p. 238, pi. 14, figs. 7, 8,9. Chamberlin, 1924, ibid., 4th ser.,
vol. 12, p. 640. Banks et al., 1932, Publ. Univ. Oklahoma Biol. Surv.,
vol. 4, no. 1, p. 22. Not Lithyphantes pulcher Keyserling.

Lithyphantes venusta Marx, 1898, in Banks, Proc. California Acad. Sci.,
3rd. ser. vol. 1, p. 239, pi. 14, fig. 2,9- Petrunkevitch, 1911, Bull.
Amer. Mus. Nat. Hist., vol. 29, p. 183 ; Roewer, 1942, Katalog der
Araneae, vol. 1, p. 408. NEW SYNONYMY.

Teutana nesiotes Chamberlin, 1924, Proc. California Acad. Sci., 4th ser.,
vol. 12, p. 639, fig. 80,9. Roewer, 1942, Katalog der Araneae, vol. 1,
p. 417. NEW SYNONYMY.

392

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Types. Female lectotype of Lithyphantes fulvus from Spring
Lake, Sevier County, Utah, designated by Fox, 1940, in the
United States National Museum. Three male and one female
syntypes of Lithyphantes parvula from San Jose del Cabo, Baja
California, in the Museum of Comparative Zoology. Female type
of Lithyphantes venusta from San Francisquito, Baja California,
destroyed. Female holotype of Teutana nesiotes from Granite
Island, north of Angel de la Guarda Island, Gulf of California,
in the California Academy of Sciences; one paratype in the
Museum of Comparative Zoology.

fP"--n-!-.-L..

u u,

Map 9. Distribution of Steatoda fulva (Keyserling).

Description. Epigynum rather small and indistinct (Figs.
46, 47), male palpus illustrated by Figures 32 and 33. Total
length of females, 3.0-5.9 mm. A female from California meas-
ured : total length 4.7 mm. Carapace 2.1 mm. long, 1.6 mm.
wide. First patella and tibia, 2.2 mm. ; second, 1.8 mm. ; third,
1.5 mm. Fourth femur, 2.0 mm. ; patella and tibia, 2.3 mm. ;
metatarsus, 1.5 mm. ; tarsiLS, 0.8 mm. Total length of males.

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 393

2.4-5.0 mm. A male from California measured : total length
5.0 mm. Carapace 2.3 mm. long, 1.9 mm. wide. First patella and
tibia, 2.8 mm. ; second, 2.4 mm. ; third, 2.1 mm. Fourth femur,
2.7 mm. ; patella and tibia, 3.1 mm. ; metatarsus, 2.3 mm. ; tarsus,
0.9 mm.

Natural History. This species is probably found under stones
in dry areas. It has also been collected in a house in Yuma,
Arizona.

Distribution. Southern and western United States south to
central Mexico. Map 9.

Records. Georgia: (Fox, 1940; Chamberlin and Ivie, 1944).
Florida: (Banks, 1904); Alachua Co.; Lake Co.; Putnam Co.
Alabama: (Banks, 1900). Louisiana: (Banks, 1899). Nebraska:
(Worley and Pickwell, 1931); Platte Co.; Columbus, June 26,
1946 (M. H. Muma). OMahoma: (Banks et al., 1932). Texas:
(Fox, 1940); Brewster Co.; Dallam Co.; El Paso; Hidalgo
Co. ; Howard Co. ; Hudspeth Co. ; Llano Co. Somervell Co. ;
Starr Co. Colorado: Mesa Co. New Mexico: Hidalgo Co.
Idaho: Boise, June 15, 1941 (B. Malkin). Utah: Carbon Co.;
Salt Lake City ; Sevier Co. ; Tooele Co. ; Weber Co. Nevada: Nye
Co. Arizona: Cochise Co.; Coconino Co.; Maricopa Co.; Organ
Pipe Cactus Natl. Mon. ; Pima Co. ; Pinal Co. ; Santa Cruz Co. :
Yuma Co. Oregon: Lake Co.; Umatilla. California: Imperial
Co. ; Inyo Co. ; Los Angeles Co. ; Mono Co. ; Riverside Co. ; San
Bernardino Co. ; San Diego Co. ; Ventura Co.

Steatoda pulcher (Keyserling), new combination
Figures 37-41, 48-51 ; Map 10.

Llthyphantes pulcher Keyserling, 1882, Die Spinnen Amerikas, vol. 2, pt.
1, p. 137, pi. 6, fig. 85,9. Marx, 1898, Proc. U.S. Natl. Mus., vol. 12,
p. 522. Banks, 1910, Bull. U.S. Natl. Mus., no. 72, p. 21 (in part).
Petrunkevitch, 1911, Bull. Anier. Mus. Nat. Hist., vol. 29, p. 183 (in
part). Roewer, 1942, Katalog der Araneae, vol. 1, p. 408 (in part).
Type. Female holotype from Washington Territory in the
Museum d'Histoire Naturelle in Paris.

Description. Epigyna illustrated by Figures 49, 50; male
palpus by Figures 37-41. Total length of females 3.0-6.8 mm.
A female from Coahuila measured: total length 5.0 mm. Cara-
pace 1.8 mm. long, 1.5 mm. wide. First femur 2.2 mm.; patella

394

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

and tibia, 2.6 mm.; metatarsus, 1.8 mm.; tarsus, 0.86 mm. Sec-
ond patella and tibia 2.0 mm.; third, 1.7 mm.; fourth, 2.6 mm.
Total length of males 2.8-5.8 mm. A male from Coahuila meas-
ured: total length, 3.2 mm. Carapace, 1.4 mm. long, 1.1 mm.
wide. First femur, 1.7 mm.; patella and tibia, 2.1 mm.; meta-
tarsus, 1.5 mm. ; tarsus, 0.7 mm. Second patella and tibia 1.6
mm.; third, 1.2 mm.; fourth, 2.0 mm.

Variation. Some specimens have the fourth leg longer than
the first, in others the first is longest. There is considerable
individual variation in the shape of the epigynum (Figs. 49,

Map 10. Distribution of Steatoda pulcher (Keyserling).

50). All the largest specimens came from northern California
and Oregon, most of the smallest from Texas; specimens from
other regions are intermediate in size. The largest specimens
from northern California and Oregon are also the lightest in
color. In specimens from this region, the abdomen is usually
white with black spots, as in eastern S. albomaculata (Fig. 65)

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 395

and the venter is as in S. alhomaculata from the Rocky Moun-
tains (Fig. 64). The smallest specimens from Texas are also
often the darkest.

The male palpus shows a surprising amount of variation;
however, the cline of variation here does not agree with the
clines for color and size. The palpus of males from northern
California (Fig. 38) is very similar to that of -S^. fulva; however,
it has a larger white membrane on the ectal side of the palp at
the inner side near the base of the embolus ; the membrane covers
more than one half the circle made by the embolus. Males from
Arizona, Texas and Coahuila have a wide palpus (Figs. 39, 40).
Unfortunately we do not have males from southern California
or eastern Arizona, but they are expected to be intermediate
in this character. Arizona and Texas specimens still have the
large white membrane on the base of the embolus (Fig. 39), but
this structure becomes smaller farther south (Fig. 40). Males
from central and southern Mexico have a narrower palpus. The
white membrane, however, is smaller (Figs. 40, 41), unlike that
of the narrow California palpus. In Mexico the palpus still
differs from that of S. fulva in that the embolus of S. pulcher
describes an ellipse wider than high (Figs. 40, 41), while that
of 8. fulva is higher than wide.

Natural History. This species has been collected under rocks
in desert in Oregon.

Distribution. Pacific coast states, southwest to Oaxaca. Map
10. In several papers Mello-Leitao refers to this species as hav-
ing been found in South America. I assume he made a mistake
in identification.

Records. Texas: Big Bend Natl. Pk. ; Concho Co.; Kendall
Co.; McCulloch Co.; Terrell Co. Colorado: Archuleta Co.;
Piedra, 7000 ft. (H. and L. Levi). New Mexico: Otero Co. Ari-
zona: Cochise Co.; Grand Canyon Natl. Pk. ; Navajo Co.; Pima
Co. Oregon: Deschutes Co.: 5 mi. S. of Redmond; Jackson Co.;
Klamath Co. ; Lake of the Woods ; Lake Co. ; Lake Albert. Cali-
fornia: Eldorado Co.; Contra Costa Co.; Kings Canyon Natl.
Pk. ; Modoc Co.; Mono Co.; Plumas Co.; Riverside Co.; San
Bernardino Co.; Shasta Co.; Sierra Co.; Siskiyou Co.; Tulare
Co. Coahuila: 5 mi. W. of Saltillo; 20 mi. E. of San Pedro.
Chihuahua: Santa Barbara; Primavera, 5500-6000 ft. ; Matachic;

396 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

20 mi. W. of Matachic. Durango: Las Puentes, 700 ft.; Otinapa,
8200 ft. Jalisco: 20 mi. N. of La Qiiemada ; Ajijic; Plan de
Barrancas; west side of Lake Saynla; nr. Tequila. Hidalgo: 5
mi. S. of Zimapan. Michoacan: Tzararacua Falls nr. Uruapan.
Morelos: Acatlipa; Cocoyoc; Alpoyeca; 10 mi. S. of Temixco.
Coajomulco; Cuernavaca; 25 km. S. Cuernavaca. Puehla: At-
lixco ; 5 mi. N. of Tehuacan ; Tehuacan ; 12 mi. N. Acatlan ; 7 mi.
S. Tlacotepec ; Tlacotepec. Veracruz: Perote. Guerrero: Hoajo-
jutla; Taxco; Mexcala. Oaxaca: Oaxaca (many collections);
Monte Alban.

Steatoda albomaculata (De Geer), new combination
Figures 56-65; Map 11.

Aranea albomaculata De Geer, 1778, Memoires pour servir a ] 'histoire des
Insectes, vol. 7, p. 257, pi. 15, figs. 2-4,9.

Eudiaria corollata, C.L. Koch, 1837, Ubersicht des Arachnidensystems, vol.
1, p. 8.

Asagena corollata, C.L. Koch, 1840, in Fiiriirohr, Die Fauna von Eegens-
burg, p. 401.

Steatoda corollata, Thorell, 1856, Recensio Critica Aranearum, p. 85;
Emerton, 1882, Trans. Connecticut Acad. Sci., vol. 6, p. 21, pi. 4, fig.
5,9; 1902, The common spiders, p. 121, fig. 285,9; 1924, Canadian
Ent.. vol. 56, p. 124.

Litliyphantes corollatiis, Thorell, 1809, On European spiders, p. 94. Emer-
ton, 1876, Psyche, vol. 1, p. 130; 1877, Proc. Boston Soc. Nat. Hist.,
vol. 19, p. 70. Thorell, 1877, Bull. U.S. Geol. Surv. vol. 3, no. 2, p. 47.
Keyserling, 1884, Die Spinnen Amerikas, vol. 2, pt. 1, p. 129, pi. 6,
fig. 81,9,5. Marx, 1889, Proc. U.S. Natl. Mus.. vol. 12, p. 522. Banks,
1901, Proc. Acad. Nat. Sci., Philadelphia, 1901, p. 579; 1895a, Jour.
New York Ent. Soc, vol. 3, p. 84; 1895b, Ann. New York Acad. Sci.,
vol. 8, p. 432; 1902, Proc. U.S. Natl. Mus., vol. 25, p. 214. Bryant,
1908, Occas. Papers Boston Soc. Nat. Hist., vol. 7, p. 17. Banks, 1910.
Bull. U.S. Natl. Mus., no. 72, p. 21. Petrunkevitch, 1911, Bull. Amer.
Mus. Nat. Hist., vol. 29, p. 182. Comstock, 1912, The spider book,
p. 362, fig. 367,9. Emerton, 1920, Trans. Eoy. Canadian Inst., vol. 12.
p. 312. Crosby and Bishop, 1928, Mem. Cornell Agr. Exp. Sta., no.
101, p. 1039. Chamberlin, 1928, Canadian Ent., vol. 60, p. 93. Worley
and Pickwell, 1931, Univ. Nebraska Studies, vol. 27, p. 27. Worley,

1932, Publ. Univ. Washington Biol., vol. 1, p. 25. Chickering, 1932,
Papers Michigan Acad. Sci., vol. 15, p. 351. Chamberlin and Ivie,

1933, Bull. Univ. Utah, biol. ser., vol. 2, p. 8. Gertsch, 1935, Amer. Mus.

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA

397

Novitates, no. 792, p. 21. Comstock, 1940, The spider book, rev. ed.,
p. 377, fig. 367,9. Fox, 1940, Proc. Biol. Soc. Washington, vol. 53,
p. 41. Probably not Aranea corollaia Linnaeus, 1758.
Lithyphantes alhomacnlatns, Simon, 1914, Arachnides de Prance, vol. 6, p.
282. Wiehle, 1937. in Dahl, Die Tierwelt Deutschlands, pt. 33, p. 200,
figs. 222-227, 9, cJ. Gertsc-h, 1939, Amer. Mus. Novitates, no. 1032, p.
4. Eoewer, 1942, Katalog dcr Aranaea, vol. 1, p. 405. Lowrie, 1942,

• \ (-^—- i.i? I I );—-■--

\ / \ \ -^

^■■■^

S. albomaculata

£.

Map 11. American distribution of Steatoda aliomaculata (De Geer).

Bull. Chicago Acad. Sci., vol. 6, p. 169; 1948, Ecology, vol. 29, p. 338.
Kaston, 1948, Bull. Connecticut Geol. Nat. Hist. Surv., no. 70, p.
78, figs. 47-50,9,5 . Kurata, 1949, Canadian Ent., vol. 81, p. 127. Levi
and Levi, 1951, Zoologica, vol. 36, p. 220. Kaston, 1953, How to know
the spiders, p. 163, fig. 407,9. Locket and Millidge, 1953, British
spiders, vol. 2, p. 55, fig. 37, 9,c5. Levi and Field, 1954, Amer. Mid-

398 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

land Nat., vol. .31, p. 444. Kaston, 1955, Trans. Illinois Acad. Sci.,
vol. 47, p. 166.

Types. The type came from the shore of the Baltic Sea.

Description. Carapace, sternum, legs, dark brown. Abdomen
black often with a white line along anterior edge, venter black
with white marks (Fig. 64). Eastern specimens may have abdo-
men white or white with black marks, sometimes in the form
of a series of anterior pointing triangles making a median dorsal
black line. Eyes snbequal in size, laterals usually slightly sep-
arated from each other. The endites of the male have tubercles.
Epigynum illustrated by Figure 58, male palpus by Figures
59-61. Total length of females, 4.0-8.0 mm. A female from
Colorado measured : total length, 6.1 mm. Carapace 2.0 mm.
long, 1.8 mm. wide. First femur, 2.6 mm.; patella and tibia, 2.7
mm.; metatarsus, 2.1 mm.; tarsus, 1.0 mm. Second patella and
tibia, 2.2 mm. ; third, 1.8 mm. ; fourth, 2.6 mm. Total length of
males, 4.3-6.8 mm. A male from Colorado measured : total length,

4.8 mm. Carapace 2.2 mm. long, 1.6 mm. wide. First femur,

2.9 mm. ; patella and tibia, 3.5 mm. ; metatarsus, 2.6 mm. ; tarsus
1.1 mm. Second patella and tibia 2.6 mm.; third, 2.0 mm.;
fourth, 3.2 mm.

Variation. The coloration in eastern specimens is light (Fig.
65) while that of western ones is usually dark, the abdomen of
the latter usually being black with only a white line along the
anterior edge and a ventral white mark. The shape of the
epigynum is quite variable. The median lobe may be much wider
than long or longer than wide. The tip of the projecting radix
of the palpus varies in shape. However these differences are
apparently not geographic.

Natural History. This species is very common in the Colorado
mountains where it is found under stones in dry, open, areas
up to timberline and down to the foothills where Latrodcctus
replaces it in similar habitats. Several spiders with numerous
egg sacs are found together, along with the remains of insects,
often ants. Kaston (1948) records egg sacs to be 4.5-8.0 mm.
in diameter containing 20-33 tan eggs. In Europe this species
is also black in color in the mountains, although black individuals
may also be found in other areas (Wiehle, 1937).

Distribution. Southern England, Europe, North Africa, Si-
beria, China, Kamchatka (Wiehle, 1937). Common in western

LEVI : SPIDER GENERA CKUSTULINA AND STEATODA 399

states and northeastern states. South to central Mexico, north
to Great Slave Lake; South America. Map 11. Judging by its
wide distribution in North America and its color variation in
different parts of the range, this species is native to North
America as well as Europe.

Records. Quebec: (Emerton, 1924). Manitoba: (Emerton,
1920). Northwest Territories: (Kurata, 1949). Alberta: (Emer-
ton, 1920); Medicine Hat; Lake Athabasca. British Columbia:
(Emerton, 1920); Cascade; Mt. Arrowsmith. New Hampshire:
Cheshire Co. Massachusetts: Barnstable Co.; Dukes Co.; Essex
Co. Connecticut: New London Co. (Kaston, 1948). New York:
(Crosby and Bishop, 1928) ; Nassau Co.; Suffolk Co. Michigan:
(Chickering, 1932; Lowrie, 1948) ; Oceana Co. Indiana: (Low-
rie, 1942, 1948). Wisconsin: (Levi and Field, 1954) ; Dane Co.;
low^a Co.; Sauk Co. Illinois: (Kaston, 1955); Kankakee Co.
Minnesota: Hennepin Co. Iowa: Ames; Sioux City. South Da-
kota: Brookings; Custer Co. Nebraska: (Worley and Pickwell,
1931) ; Boone Co.; Scotts Bluff Co. Montana: (Gertsch, 1939) ;
Helena. Wyoming: (Levi and Levi, 1951); Cheyenne; Goshen
Co. ; Lincoln Co. ; Niobara Co. ; Teton Co. ; Washakie Co. Col-
orado: (Banks, 1895b; Fox, 1940); Archuleta Co.; Boulder
Co. ; Chaffee Co. ; El Paso Co. ; Gilpin Co. ; Gunnison Co. ;
Hinsdale Co. ; Jefferson Co. ; Lake Co. ; Larimer Co. ; Mineral
Co. ; Rio Grande Co. ; Routt Co. ; Teller Co. New Mexico: Catron
Co. ; Otero Co. ; San Miguel Co. Idaho: Bannock Co. ; Bear Lake
Co.; Bingham Co.; Fremont Co.; Power Co. Utah: (Chamberlin
and I vie, 1933 ; Fox, 1940) ; Bryce Canyon Natl. Pk. ; Kane Co. ;
Millard Co.; San Juan Co.; Sevier Co. Arizona: (Banks, 1902) ;
Flagstaff ; Grand Canyon Natl. Pk. ; Navajo Co. ; Pima Co.
Washington: (Worley, 1932) ; Mt. Ranier Natl. Pk. ; San Juan
Co.; Stevens Co.; Walla Walla; Whitman Co. Oregon: Baker
Co. ; Deschutes Co. ; Harney Co. ; Hood River Co. ; Jackson Co. ;
Umatilla Co.; Wallowa Co.; Wheeler Co. California: Alpine
Co. ; Eldorado Co. ; Inyo Co. ; Lassen Co. ; Los Angeles Co. ;
Modoc Co. ; Mono Co. ; Nevada Co. ; Placer Co. ; Sacramento Co. ;
Siskiyou Co. ; Yosemite Natl. Pk. ; Yuba Co. Chihuahua: Los
Canoas Babicora; 7700 ft. summit NE of San Jose Babicora;
Matachic. San Luis Potosi: 12 mi. W. of Arriaga.

400 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

Steatoda AMERICANA (Emerton), new combination
Figures 66-69 ; Map 12.

Asar/ena americana Emerton, 1882, Trans. Connecticut Acad. Sci., vol. 6,
p. 23, pi. 4, fig. 6,S. Keyserling, 1886, Die Rpinnen Amerikas, vol. 2.
pt. 2, p. 2 pi. 11, fig. 135,5. Marx, 1889, Proc. U.S. Natl. Museum,
vol. 12, p. 525. Simon, 1894, Histoire naturclle des Araignees vol. 1,
p. 574, fig. 586. Banks, 1895, Jour. New York Ent. Soc, vol. 3, p. 84;
1895, Ent. News, vol. 6, p. 205. Emerton, 1902, The common spiders,
p. 122, fig. 287,9. Banks, 1904, Proc. Acad. Nat. Sci. Philadelphia,
vol. 56, p. 653; 1907, Indiana Dept. Geol. Nat. Eesources, 31st Rept.,
p. 739; 1908, Proc. Ent. Soc. Washington, vol. 9, p. 5; 1910, Bull. U.S.
Natl. Mus., no. 72, p. 22; 1911, Proc. Acad. Nat. Sci. Philadelphia, vol.
63, p. 445. Petrunkevitch, 1911, Bull. Amer. Mus. Nat. Hist., vol. 29,
p. 170. Comstock, 1912, The spider book, p. 364, fig. 369,$. Emerton,
1913, Appalachia, vol. 12, p. 155. Barrows, 1918, Ohio Jour. Sci., vol.
18, p. 302. Emerton, 1920, Trans. Roy. Canadian Inst., vol. 12, p.
312. Bishop, 1925, Bull. New York State Mus., no. 260, p. 66, fig. 1,^.
Bishop and Crosby, 1926, Jour. Elisha Mitchell Sci. Soc, vol. 41, p.
176. Crosby and Bishop, 1928, Mem. Cornell Agr. Exp. Sta., no. 101,
p. 1039. Chamberlin, 1928, Proc. Biol. Soc. Washington, vol. 41, p
179. Worley and Pickwell, 1931, Univ. Nebraska Studies, vol. 27, p
25. Worley, 1932, Publ. Univ. Washington Biology, vol. 1, p. 23
Banks, Newi^ort and Bird, 1932, Publ. Univ. Oklahoma, Biol. Surv., vol
4, no. 1, p. 21. Chickering, 1934, Papers Michigan Acad. Sci., vol. 19
p. 578. Fox, 1940, Proc. Biol. Soc. Washington, vol. 53 p. 40. Com-
stock, 1940, The spider book, rev. ed., p. 379, fig. 369,9. Chamberlin
and Ivie, 1944, Bull. Univ. Utah, biol. ser., vol. 8, no. 5, p. 37. Muma,
1945, Bull. Univ. Maryland Agr. Exp. Sta., no. A38, p. 24. Muma and
Jeffers, 1945, Ann. Ent. Soc. America, vol. 38, p. 248. Archer, 1946,
Paper Alabama Mus. Nat. Hist., no. 22, p. 22. Levi and Field, 1954.
Amer. Midland Nat., vol. 51, p. 442. Kaston, 1955, Trans. Illinois
Acad. Sci., vol. 47, p. 166. Kaston, 1948, Bull. Connecticut Geol. Nat.
Hist. Surv., no. 70, p. 73, figs. 25-29, $,c5. Kaston, 1953, How to
know the spiders, p. 163, fig. 406, $ . Elliott, 1953, Proc. Indiana
Acad. Sci., vol. 62, p. 308. Bonnet, 1955, Bibliographia Araneorum,
vol. 2, p. 753.

Asagena dubia, Roewer, 1942, Katalog der Araneae, vol. 1, p. 397. Probably
not Herpyllus dubia Hentz.

Types. Male holotype from garden fence in Boston, Mass.,
May 30, in the Museum of Comparative Zoology.

Description. Carapace, sternum brown ; legs yellow-brown.
Abdomen black, often with a pair of white spots on posterior

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA

401

portion of dorsum. Anterior median eyes sometimes slightly
smaller than others, almost their diameter apart, three-quarters
diameter from laterals. Posterior eyes one diameter apart, one
diameter from laterals. Tooth on anterior margin of male
chelicerae indistinct. Males with femora swollen, often with
ventral spines on first femora and tibia, but these may be absent.
Usually with a thorn on distal end of second femur. Epigynum
illustrated by Figure 67, male palpus by Figure 68. Total length
of females 3.5-4.7 mm. A female from Wisconsin measured:
total length, 4.0 mm. Carapace 1.6 mm. long, 1.3 mm. wide.

if?=^

S omericano

Map 12. Distribution of Steatoda americana (Emerton).

First patella and tibia, 1.5 mm.; second, 1.2 mm.; third 1.0 mm.
Fourth femur, 1.2 mm. ; patella and tibia, 1.6 mm. ; metatarsus,
0.7 mm. ; tarsus, 0.6 mm. Total length of males, 3.2-4.4 mm. A
male from Wisconsin measured : total length, 4.4 mm. Carapace
2.2 mm. long, 1.6 mm. wide. First patella and tibia, 1.7 mm. ;
second, 1.2 mm. ; third, 1.4 mm. Fourth femur, 1.5 mm. ; patella
and tibia, 1.7 mm. ; metatarsus, 1.2 mm. ; tarsus, 0.7 mm.

Natural History. This species is found in leaf litter, under
stones, under loose bark of logs and in moss; it has also been
found in Wisconsin sweeping grass and vegetation in woods. It
has been found with three spherical, transparent, egg sacs, 4-5

402 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

mm. in diameter, containing 22-33 light yellow eggs (Kaston,
1948). Muma and Jeffers (1945) have found it in nests of the
wasp Sceliphron. It has also been reported from other wasp nest
collections.

Distribution. United States and southern Canada, except Cali-
fornia and Oregon, south to Hidalgo.

Records. Ontario: Rockport. British Columbia: (Emerton,
1920). Maine: Cumberland Co. New Hampshire: (Emerton,
1913). Vermont: Chittenden Co. Rhode Island: (Fox, 1940).
Connecticut: (Kaston, 1948). New York: (Crosby and Bishop,
1928) ; Bronx; Onondaga Co.; Suffolk Co. New Jersey: Bergen
Co. Maryland: (Muma, 1945). District of Columbia: Washing-
ton. Virginia: Fairfax Co. West Virginia: (Bishop and Crosby,
1924); Pocahontas Co. North Carolina: (Banks, 1911; Bishop
and Crosby, 1926) ; Swain Co. Georgia: (Chamberlin and Ivie,
1944). Florida: (Banks, 1904). Alabama: (Archer, 1946).
Ohio: (Barrows, 1918) ; Columbiana Co.; Cuyahoga Co. Michi-
gan: (Chickering, 1934). Indiana: (Elliott, 1953); Porter Co.
Tennessee: Roane Co. Wisconsin: (Levi and Field, 1954) ; Craw-
ford Co.; Door Co.; Grant Co.; Richland Co.; Vernon Co.
Illinois: (Kaston, 1955) ; Randolph Co. Missouri: (Banks,
1895) ; Crawford Co.; St. Louis. Nebraska: (Worley and Pick-
well, 1931). Oklahoma: Cleveland Co. Texas: San Antonio.
Colorado: Archuleta Co.; Denver. New Mexico: Catron Co.;
Otero Co. (9-12000 ft.). Idaho: Bear Lk. Co. Utah: (Chamber-
lin, 1928); Salt Lake City; Zion Natl. Pk. Arizona: Mohave
Co. Washington: (Worley, 1932). Oregon: Salem. Sonora:
27 mi. S. of Nogales. Hidalgo: S. of Jacala, July 20, 1956 (V.
Roth, W. J. Gertsch).

Steatoda septemmaculata (Keyserling), new combination
Figures 70-73; Map 13.

Lithyplmntes septemmaculatus Keyserling, 1884, Die Spiunen Amerikas, vol.
2, no. 1, p. 141, pi. 6, fig. 88,9. van Hasselt, 1887, Tijdschr. Ent., vol.
30, p. 234. Marx, 1889 Proc. U.S. Natl. Mus., vol. 12, p. 522. Banks,
1904, Proc. Acad. Nat. Sci. Philadelphia, vol. 56, p. 653; 1910, Bull.
U.S. Natl. Mus. uo. 72, p. 21. Petrunkevitch, 1911, Bull Amer. Mus.
Nat. Hist., vol. 29, p. 183; 1930, Trans. Connecticut Acad. Sci., vol.
30, p. 169, figs. 6, 7,9. Bryant, 1940, Bull. Mus. Comp. Zool., vol.

LEVI : SPIDER GENERA CBUSTULINA AND STEATODA

408

86, p. 304. Eoewer, 1942, Katalog der Araneae, vol. 1, p. 408. Bryant,
1942, Bull. Mus. Comp. Zool., vol. 89, p. 337; 1945, Trans. Connecticut
Acad. Sei., vol. 36, p. 204, pi. 1, fig. 7, 9, S ; 1948, Bull. Mus. Comp.
Zool., vol. 100, p. 375. ? Kaston, 1948, Bull. Connecticut Geol. Nat.
Hist. Surv., no. 70, p. 79.
Types. Sj'iitypes from "Denver in Columbia" collected bj-

Marx, and also from Enterprise, Florida, in the United States

National Museum. Marx's labels are commonly wrong and the

Denver locality is undoubtedly an error.

Description. Carapace, sternum, legs orange. Femora dusky.

Abdomen purple-black with seven white spots on dorsum (Fig.

70), two of them above spinnerets. Eyes subequal in size. An-

S septemmoculoto

^ .cM^ »*-• '

Map 13. Distribution of Steatoda septemmaculata (Keyserling).

terior median eyes one diameter apart, less than one diameter
from laterals. Posterior medians one diameter apart, one and
one-half diameters from laterals. Laterals slightly separated
from each other. Chelicerae and maxillae of male modified; the
former lacking teeth. Epigynum illustrated by Figure 72, male
palpus by Figure 73. Total length of females 2.5-3.2 mm. A
female from Florida measured total length 2.5 mm. Carapace
1.10 mm. long, 0.88 mm. wide. First patella and tibia, 1.00 mm. ;
second, 0.89 mm. ; third, 0.75 mm. Fourth femur, 1.04 mm. ;

404 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

patella and tibia, 1.18 mm.; metatarsus, 0.62 mm.; tarsus, 0.47
mm. Total length of males 1.8-3.4 mm. A male from Florida
measured total length, 3.0 mm. Carapace 1.78 mm. long, 1.11
mm. wide. First patella and tibia, 1.26 mm.; second 1.15 mm.;
third, 0.98 mm. Fourth femur, 1.28 mm.; patella and tibia,
1.35 mm.; metatarsus, 0.84 mm.; tarsus, 0.60 mm.

Natural History Petrunkevitch (1930) found this species on
the undersides of old coconut shells and in cactus fields in
Puerto Rico.

Distribution. Florida, Panama, West Indies. (Map 13.) Kas-
ton (1948) reported a female found in Litchfield, Connecticut.
This has presumably been imported, or it is an erroneous record.
The specimen could not be found.

Records. Florida: (Banks, 1904; Bryant, 1945) Collier Co.;
De Soto Co.; Hillsborough Co.; Orange Co.; Polk Co. Yucatan:
Cayo Arenas. Panama: Santa Rosa (Colon) ; Ensenada. Isla de
Providencia. Cuba: (Bryant, 1940); Habana; Cabanas; Dai-
quiri; Pinar del Rio; Marianao. Jamaica: Western Kingston.
Navassa Isl. Hispaniola: (Bryant, 1948). Puerto Bico: (Pe-
trunkevitch, 1930). Virgin Islands: (Bryant, 1942). Lesser
A^iiilles: (van Hasselt, 1887).

Steatoda grossa (C.L. Koch)
Figures 74, 83-85 ; Map 14.

Theridion grossum C.L. Koch, 1838, Die Arachniden, vol. 4, p. 112, fig.

321,?.
Steatoda grossa, C.L. Koch, 1851. Ubersicht des Araelmidensystems, pt. 5,

p. 17.

Theiridion nitidum Hobiiberg, 1876, An. Agr. Argentina, vol. 4, p. 72.

Th.eridion domesticum Holiuberg, 1876, op. cit.

Steatoda pusulosa Keyseiling, 1877, Verhandl. zool. bot. Gesell. W^ien., vol.
27, p. 579, pi. 14, fig. 7, 8,?, 5.

Lithyphantes grossa, Pavesi, 1878, Ann. Mus. Civ. Geneva, vol. 11, p. 371.

Teutana grossa, Simon, 1881, Les Araehnides de France, vol. 5, p. 164
Banks, 1898, Proc. California Acad. Sci., 3rd ser., vol. 1, p. 238, F.O.P
Cambridge, 1902, Biologia Central! Americana, Araneidea, vol. 2, p
374, pi. 35, fig. 9, 10,5,0. Banks, 1909, Pvoc, Acad. Nat. Sci. PhUa
delphia, vol. 61, p. 205. Petrunkevitch, 1911, Bull. Amer. Mus. Nat
Hist., vol. 29, p. 189. Emerton, 1911, Trans. Connecticut Acad. Sci.,
voi. 16, p. 387, fig. 2,<5. Chamberlin, 1919, Jour. Ent. Zool., vol. 12,

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 405

p. 25. Beimoser, 1939, Ann. Naturhist. Mus. Wien, vol. 50, p. 344.
Pox, 1940, Proc. Biol. Soc. Washington, vol. 53, p. 42. Chamberlin and
Ivie, 1941, Bull. Univ. Utah, bid. ser., vol. 6, no. 3, p. 12. Roewer,
1942, Katalog der Araneae, vol. 1, p. 414. Branch, 1942, Bull. South.
California Acad. Sei., vol. 41, p. 138. Archer, 1946, Paper Alabama
Mus. Nat. Hist., no. 22, p. 21. Kaston, 1948, Bull. Connecticut Geol.
Nat. Hist. Surv., no. 70, p. 86, figs. G0-62,$,<5. Schenkel, 1950, Ver-
handl. Naturf. Gesell. Basel, vol. 61, p. 52.
Teutana nitida, Keyserling, 1882, Die Spinnen Amerikas, vol. 2, pt. 1, p.

124, pi. 6, fig. 79, 9, S.
Teutana sonata Keyserling, 1882, op. int., p. 127, pi. 6, fig. 80, <5. Banks,
1898, Proc. California Acad. Sci., 3rd ser. vol. 1, p. 238. O.P. Cam-
bridge, 1902, Biologia Centrali Americana, Araneidea, vol. 1, p. 306,
pi. 35, fig. 7, 8,9,S. NEW SYNONYMY.
Asag.ena sonata, F.O.P., Cambridge, 1902, ibid, vol. 2, p. 378. Petrunkevitch,
1911, Bull. Amer. Mus. Nat. Hist., vol. 29, p. 171. Roewer, 1942,
Katalog der Araneae, vol. 1, p. 397, Bonnet, 1955, Bibliographia
Araneorum, vol. 2, p. 757. NEW SYNONYMY.
Teutana modesta Bryant, 1948, Bull. Mus. Comp. Zool., vol. 100, p. 375,
figs. 49, 51,9. NEW SYNONYMY.
Types. The types of Theridion grossum from Peloponnesus
peninsula and Germany. The types of Theridion nitidum and
Theridion domesticum Holmberg from Argentina are lost. The
types of Steatoda pusulosa Keyserling from Uruguay. Male
type of Teutana zona fa Keyserling from Mexico in the Museum
National Histoire Naturelle in Paris. Female holotype of Teu-
tana modesta Bryant from the Cordillera Central, Valle Nuevo.
7000 ft., Dominican Republic in the Museum of Comparative
Zoology.

Description. Carapace, sternum, legs dark orange. Abdomen
purple-black, with an indistinct light line around anterior mar-
gin and several indistinct light patches on dorsum or with pat-
tern as in S. triangulosa. Eyes subequal in size. Anterior eyes
separated by one-third their diameter. Posterior medians sep-
arated by one-half their diameter, by one diameter from laterals.
Chelicerae of males swollen, with a short thick fang. Epigynum
variable, sometimes with a wide septum (Fig. 84) sometimes with
septum almost missing (Fig. 85). Male palpus illustrated by
Figure 74. Total length of females 5.9-10.5 mm. A female from
California measured : total length, 8.6 mm. Carapace 3.2 mm.
long, 2.7 mm. wide. First femur, 4.5 mm. ; patella and tibia,

406

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

5.6 mm. ; metatarsus, 4.3 mm. ; tarsus, 1.9 mm. Second patella
and tibia, 4.0 mm.; third, 3.2 mm., fourth, 4.5 mm. Total length
of males 4.1-7.2 mm. A male from California measured : total
length, 6.5 mm. Carapace 3.1 mm. long, 2.3 mm. wide. First
femur, 4.7 mm. ; patella and tibia, 5.5 mm. ; metatarsus, 4.6 mm. ;
tarsus, 1.8 mm. Second patella and tibia, 4.1 mm.; third, 3.2
mm. ; fourth, 4.7 mm.

^.

^) — :l ;

•V/ ; \ J I \

S grosso

^=?==^2^iiP"

Map 14. North and Central American distribution of Steatoda grossa
(C.L. Koch).

Natural History. Branch (1942) raised /S. grossa in the lab-
oratory, feeding it on Drosophila and house flies. This species
seems to be common in cities in North America. Archer (1946)
reports that it lives in brick piles and under loose bricks and
that it preys on Latrodectus mactans (Fabricius), the black-
widow spider.

Distribution. Cosmopolitan. In the United States, along the
coasts only (Map 14). The proximity of this species to man
would suggest that it has been introduced recently into North
America.

Records. Massachusetts: (Emerton, 1911). Rhode Island:
Providence. Connecticut: (Kaston, 1948). Georgia: Daugherty

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 407

Co. Florida: Alachua Co. Alabama: Mobile; (Archer, 1946).
Mississippi: Forrest Co. Louisiana: East Baton Rouge Par.;
Orleans Par.; West Baton Rouge Par. Washington: Olympia;
Pacific Co.; Seattle. Oregon: Corvallis; Curry Co.; Jackson
Co.; Eugene; Hood River Co.; Portland; Tillamook Co. Cali-
fornia: (Chamberlin and Ivie, 1944; Schenkel, 1950) ; Alameda
Co. ; Del Norte Co. ; Los Angeles Co. ; Marin Co. ; Monterey Co. ;
Mendocino Co. ; Sacramento, San Diego Co. ; San Francisco ;
San Nicolas Isl. ; Santa Barbara; Yolo Co. Chihuahua: Santa
Barbara. Baja California: Isl. Cedros; Isl. Natividad. Sonora:
(Banks, 1898). Zacatecas: 10 mi. E. of Zacatecas. Puehla:
Tehuacan. Distrito Federal: San Geronimo; San Angel; Coaya-
can; Mexico (in house). Morelos: Cuernavaca. Michoacan: Za-
mora. Veracruz: Los Naranjos. Costa Rica: (Banks, 1909; Rei-
mo.ser, 1939). Jamaica: St. Andrew Par. Peru: Arequiba; Aco-
mayo (Huanuco) ; Tingo Maria; Lima. Brazil: Minas Gerais;
Min. Serinha Diamantina. Argentinu: (Keyserling, 1886) ;
Salta.

Steatoda triangulosa (Walckenaer)
Figures 75, 76, 80-82; Map 15.

Aranea triangulosa Walckenaer, 1892, Faune Parisienne, vol. 2, p. 207.
Theridion triangulifer, Walckenaer, 189.5, Tableau des Araneides, p. 75, pi.

8, fig. 73, 74.
Theridion serpentinum Hentz, 1850, Jour. Boston Soc. Nat. Hist., vol. 6,

p. 273, pi. 9, fig. 2; 1875, The spiders of the United States, p. 144, pi.

16, fig. 2.
Steatoda trianguUfera, Simon, 1873, Mem. Soc. Roy. Sci. Li&ge, vol. 5, p.

116.
Steatoda triangulosa, Thorell, 1873, Remarks on synonyms of European

spiders, p. 505. Emerton, 1882, Trans. Connecticut Acad. Sci., vol. 6,

p. 22, fig. 6,5. Banks, 1892, Proc. Acad. Nat. Sci. Philadelphia, p.

31. Emerton, 1902, The common spiders, p. 121, fig. 286,$.
Teutana triangulosa, Simon, 1881, Les Arachnides de France, vol. 5, p. 163.

Keyserling, 1884, Die Spinnen Amerikas, vol. 2, pt. 1, p. 122, pi. 6, fig.

78, $,5. Marx, 1889, Proc. U.S. Natl. Mus., vol. 12, p. 521. Banks,

1895, Ent. News, vol. 6, p. 205; 1896, Jour. New York Ent, Soc, vol.

4, p. 191; 1899, Proc. Ent. Soc. Washington, vol. 4, p. 189; 1900, Proc.

Acad. Nat. Sci. Philadelphia, p. 533. Montgomery, 1903, ibid., p. 112.

Scheffer, 1905, Trans. Kansas Acad. Sci., vol. 19, p. 192. Bryant,

408 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1908, Occas. Papers Boston Soc. Nat. Hist., vol, 7, p. 17. Banks, 1910,
Bull. U.S. Natl. Mus., no. 72, p. 21; 1911, Proc. Acad. Nat. Sci.
Philadelphia, vol. 63, p. 445. Petrunkevitch, 1911, Bull. Amer. Mus.
Nat. Hist., vol. 29, p. 190. Comstock, 1912, The spider book, p. 361,
figs. 364, 365,9. Barrows, 1918, Ohio Jour. Sci., vol. 18, p. 304. Bishop
and Crosby, 1926, Jour. Elisha Mitchell Sci. Soc, vol. 41, p. 181,
Crosby and Bishop, 1928, Mem. Cornell Agr. Exp. Sta., no. 101, p. 1041.
Worley and Pickwell, 1931, Univ. Nebraska Studies, vol. 27, p. 29.
Banks, Newport and Bird, 1932, Publ. Univ. Oklahoma, Biol. Surv.,
vol. 4, no. 1, p. 22. Wiehle, 1937, in Dahl, Die Tierwelt Deutschlands,
pt. 33, p. 198, figs. 217-221, 9, (J. Stiles and Detwiler, 1938, Proc. Iowa
Acad. Sci., vol. 45, p. 286. Comstock, 1940, The spider book, rev. ed.,
p. 376, figs. 364-365,9. Fox, 1940, Proc. Biol. Soc. Washington, vol.
53, p. 42. Roewer, 1942, Katalog der Araneae, vol. 1 p. 416. Chamber-
lin and Ivie, 1944, Bull. Univ. Utah, biol. ser., vol. 8, no. 5, p. 46.
Muma, 1945, Bull. Univ. Maryland Agr. Exp. Sta., no. A38, p. 27.
Archer, 1946, Paper Alabama Mus. Nat. Hist., no. 22, p. 21. Kaston,
1948, Bull. Connecticut Geol. Nat. Hist. Surv., no. 70, p. 86, figs.
57-59, 9 ,$ . Kaston, 1953, How to know the spiders, p. 165, fig. 413, 9 .
Elliott, 1953, Proc. Indiana Acad. Sci., vol. 62, p. 309. Levi and Field,
1954, Amer. Midland Nat., vol. 51, p. 444. Kaston, 1955, Trans. Illinois
Acad. Sci., vol. 47, p. 166.
Theridion saylori Fox, 1940, Proc. Biol. Soc. Washington, vol. 53, p. 43,
fig. 1,9.
Types. Types of At^anea triangulosa from Paris, France. Type
of Theridion serpentinum from Georgia is lost. Type of Ther-
idion saylori from Ozark Lake, Missouri in the United States
National Museum.

Description. Carapace, sternum orange-yellow; legs yellow.
Abdomen purplish brown with white spots on dorsum (Fig. 82)
and towards sides. Eyes subequal or anterior medians smaller.
Anterior median eyes separated by two-thirds their diameter,
by one-third to two-thirds from laterals. Posterior medians two-
thirds to one diameter apart, one diameter from laterals. Epigy-
num illustrated by Figure 81, male palpus by Figure 76. Total
length of females, 3.6-5.9 mm. A female from Massachusetts
measured : total length, 5.1 mm. Carapace 1.9 mm. long, 1.8 mm.
wide. First femur, 3.0 mm. ; patella and tibia, 3.8 mm. ; meta-
tarsus, 2.6 mm. ; tarsus, 1.3 mm. Second patella and tibia, 2.4
mm. ; third, 1.6 mm. ; fourth, 2.7 mm. Total length of males
3.5-4.7 mm. A male from Massachusetts measured : total length,
4.7 mm. Carapace 2.3 mm. long, 1.8 mm. wide. First femur

LEVI : SPIDER GENERA CKUSTULINA AND STEATODA

409

3.2 mm. ; patella and tibia, 3.7 mm. ; metatarsus, 2.4 mm. ; tarsus,
2.0 mm. Second patella and tibia, 2.5 mm.; third, 2.0 mm.;
fourth, 2.7 mm.

Natural History. This species is locally common in and on
houses. In museums it may infest dermestid beetle cultures.
Wiehle (1937) reports that it lives in houses in the northern
part of its range in Europe, under stones and on walls in the
southern part of the range. This agrees with Archer's (1946)
observations in the southern United States. Montgomery (1903)
observed mating of this species.

Map 15. North American distribution of Steatoda triangulosa (Walcke-
naer).

Distrihution. Central and southern Europe, southern Russia,
Mediterranean. St. Helena (Wiehle, 1937). United States. Map
15. South America. The city dwelling habit would suggest that
this species has been introduced to North America recently.

Records. Massachusetts: (Bryant, 1908) ; Bristol Co.; Spring-
field. Connecticut: (Kaston, 1948). Neiv York: (Banks, 1896;
Crosby and Bishop, 1928) Rochester; Bronx; Manhattan; Brook-
lyn. New Jersey: (Fox, 1940) ; Bergen Co.; Camden Co.; Mer-
cer Co. Pennsylvania: Lancaster; Northampton Co.; Philadel-
phia. Maryland: (Keyserling, 1884; Muma, 1945). District of
Columbia: (Fox, 1940). Virginia: Fairfax Co. ^Yest Virginia:
Monongahela Co. North Carolina: (Banks, 1911; Fox, 1940);
Mecklenburg Co.; Union Co. f^outh Carolina: Georgia: (Hentz,

410 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

1850; Chamberlin and Ivie, 1944). Alabama: (Archer, 1946);
Colbert Co.; Tallapoosa Co. Louisiana: (Banks, 1899). Ohio:
(Hentz, 1875; Barrows, 1918). Michigan: Washtenaw Co. In-
diana: (Elliott, 1953). Wisco7isin: (Levi and Field, 1954).
Illinois: Cook Co.; Urbana. Iowa: (Stiles et al. 1938). Missouri:
(Banks, 1895). ArJcansas: Washington Co. Nebraska: (Worley
and Pickwell, 1931). Kansas: (Scheffer, 1905); Cowley Co.
Oklahoma: (Banks et al., 1932) ; Payne Co. Texas: Atascosa
Co. ; Austin ; Dallas ; Kerr Co. ; McLennan Co. ; San Antonio.
Colorado: Boulder; Denver. Idaho: Bear Lake. Utah: Emery
Co.; Salt Lake City. Oregon: Jackson Co.; Lane Co.; Wasco Co.
California: Alameda Co.; Solano Co.

Steatoda castanea (Clerck)
Figures 77-79

Araneus castaneus Clerek, 1757, Aranei Suecici, p. 49, pi. 3, tig. 3.
Teutana castanea, Wiehle, 1937, mi Dahl, Die Tierwelt Deutschlands, pt. 33,
p. 195, figs. 205 211,9,5. Eoewer 1942, Katalog der Araneae, vol. 1.
p. 415.
One female has been collected at St. Thomas, Ontario, prob-
ably in a greenhouse. It has not become established in this
region.

Distribution. Europe. Turkestan (Wiehle, 1937).

Steatoda bipunctata group

The group is represented by a series of closely related species
with the eastern Steatoda borealis at one extreme, the European
S. bipunctata at the other. In North America they present a
series of mostly allopatric forms, each slightly different from
the preceding. The last two of these species {grandis, americana)
overlap one of the preceding (hes'pera). Furthermore, these
last two are sympatric over part of their range. An attempt was
made to cross one female 8. hespera with males of S. borealis
in the laboratory. No egg sac was produced. This, however,
should be repeated.

Description. The coloration, size, proportions of all species
are about the same. The carapace is yellow to red-brown, ster-
num and legs brown, coxae lighter brown. Dorsum of abdomen

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 411

dark brown to purplish black with a median white line and
an anterior marginal line which continues on sides (Fig. 91).
Venter light with two dark marks, or rarely all yellow or all dark
(Fig. 90). Short spines covering thorax, more distinct in male.
Carapace of males more rugose. Anterior median eyes larger
(about 1.3 diameter) than others. Anterior medians two-thirds
diameter apart, less than one-fourth diameter from laterals.
Posterior medians separated by about one diameter, slightly
closer to laterals. Laterals slightly separated from each other
in many species. Epigynum similar in all species. Internal
genitalia heavily sclerotized, not very useful in differentiating
species. Male palpus with large radix (R in Figs. 153-154). A
female Steatoda horealis from Wisconsin measured : total length,
4.8 mm. Carapace 1.8 mm. long, 1.6 mm. wide. First femur, 2.2
mm. ; patella and tibia, 2.6 mm. ; metatarsus, 1.6 mm. ; tarsus, 1.0
mm. Second patella and tibia, 1.8 mm. ; third, 1.4 mm. ; fourth,
2.2 mm. A male from Wisconsin measured : total length, 4.5 mm.
Carapace 2.0 mm. long, 1.6 mm. wide. First femur, 2.3 mm.;
patella and tibia, 2.9 mm. ; metatarsus, 1.9 mm. ; tarsus, 1.1 mm.
Second patella and tibia, 2.0 mm. ; third, 1.6 mm. ; fourth,
2.2 mm.

Variation. Three species are polymorphic (>S'. atascadera, S.
mexicana, S. grandis). The genitalia of 8. atascadera seem more
variable in that part of its range where no other species of the
S. bipunctata group are found, than in the northern part of its
range where S. hespera is also found. The male palpi of S.
grandis are quite uniform in structure, but the epigyna are ex-
tremely variable. Both the palpi and epigyna of 8. mexicana are
variable. Where collected together, 8. grandis and 8. mexicana
always are quite distinct; however females from many localities
may be difficult to place. It is assumed that in these latter local-
ities only one or the other species occurs.

The question may be raised here whether 8. horealis and 8.
hespera are two species or subspecies. There is no overlap in the
two groups and the distribution would indicate that only one
species is involved. However males and females of the two
species in the Rocky Mountain region are always easy to place
in one species or another, while some female 8. hespera from
the West Coast appear much more like 8. horealis. In other
words the species seem more distinct along the common border

412 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

of their range than in other parts of it, which would indicate
that we are dealing with two species.

Natural History. The species of Steatoda live under bark,
in crevices of rocks, sometimes under stones. They are common
in buildings wdiere they build small webs in corners ; however,
during daytime the spiders stay hidden in crevices, coming out
to the web in the evening. Females and males may inhabit the
same web. In captivity slight vibration of flies in the web, or
sometimes only the presence of a dead fly in the web, brought
the spiders out of their retreats. Kaston (1948) described the
egg sac of S. horealis as being "loosely woven, so that the ivory
to pale pink eggs show through the brownish white threads. The
shape is approximately spherical 5-6.5 mm. in diameter and I
counted in four egg sacs, 37, 47, 76, and 95 eggs respectively."

Distribution. This group may be mainly holarctic in distri-
bution. The male Steatoda hrasiliana Keyserling from Brazil
may be an introduced form.

Key to females of the Steatoda bipunctata group

Note that figures were drawn with the epigynum lying flat, thus from
a slightly anterior position of the abdomen.
]. Raised anterior median lobe of epigynum wider than long (Fig.
113) 2

1. Raised anterior median lobe as long as wide, or longer than wide (Fig.

93, 105, 109) 4

2. Anterior arms of depression pointing straight anterior (Figs. 107-109),

California coast range (Map 17) atascadera

2. Arms of depression pointing slightly to sides (Figs. Ill, 113, 115) ... .3

3. Posterior portion of depression swollen and wider than arms (Figs.

117, 118). East of Rocky Mountains to Alaska (Map 19) . . .horealis

3. Posterior portion of depression as wide as arms (Figs. Ill, 113, 115)

Western United States to southern British Columbia (Map 19) ....
hespera

4. Raised anterior median lobe as wide as long (Figs. 107, 109). Cali-

fornia coast range (Map 17) atascadera

4. Raised anterior median lobe longer than wide 5

5. Found in southern California (Map 17) palomara

5. Not found in southern California . 6

r>. With a large swollen posterior lip and swollen areas on each side of

depression (Fig. 89) bipunctata

(i. Lacking swollen areas on each side of depression 7

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 413

7. Openings in a deep depression on sides of a very narrow septum (Figs.
93-97). Depression often with material difficult or impossible to
remove. Western United States (Map 17) grandis

7. Openings in a shallower depression, septum wider (Figs. 99-103).
Material in depression easily removed. Western United States and
Mexico (Map 18) mexicana

Key to males of the Steatoda bipunctata group

Note that figures of the palpus in mesal view were drawn with the radix
lying flat, thus from a slightly anterior view.
1. Radix with a well-developed median lobe (Figs. 148, 153). East of
Rocky Mountains to Alaska (Map 19) borealis

1. Radix lacking median lobe (Figs. 119, 129, 142, 155) 2

2. Radix enclosing an area, the diameter of which is less, equal or only

slightly more than the greatest width of radix (Figs. 119, 155) . . .3

2. Radix enclosing an area, the diameter of which is much greater than

the width of the radix (Figs. 129, 145) 4

3. Embolus short and thick (Fig. 156) bipunctata

3. Embolus long and thin (Fig. 123). Western United States (Map 17)

grandis

4. Radix with a distinct ectal lobe below tip (Figs. 130-132, 135, 137, 139,

141) 5

4. Radix with no lobe near tip 6

5. California coast range south to Riverside County (Map 17)

atascadera

5. Southern California (Map 17) palomara

6. Radix with tip pointing eetally, above a notch, (Figs. 130-132)

Southern California (Map 17) palomara

6. End of radix pointing ectal but without notch below (Figs. 125,

146) 7

7. Base of embolus almost circular with rim curved outward (Figs. 144,

147) Western United States, southern British Columbia (Map 19)
hespera

7. Base of embolus oval with rim not curved outward (Fig. 128) Western
United States and Mexico (Map 17) mexicana

Steatoda bipunctata (Linnaeus)
Figures 86-89, 155-156 ; Map 16.

Aranea bipunctata Linnaeus, 1758, Systema Naturae, 10th ed., p. 620.
Aranea 4-punctata, Fabrieus, 1775, Systema Entomologiae, p. 434.
Aratnea punctata, De Geer, 1778, Memoires pour servir h. I'histoire des

Insectes, vol. 7, p. 255, pi. 15, fig. 1.
Theridion 4-punctatum, Walckenaer, 1805, Tableau des Aran6ides, p. 73, pi.

7, figs. 69, 70.

414

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Steatoda 4-punctatum, Sundevall, 1833, Conspectus Arachnidum, p. 17.

Phrurolithns ornatu.'i C.L. Koch, 1839, Die Arachniden, vol. 6, p. 114, fig
515.

Eiicharia hipunctata, C.L. Koch, 1845, ibid., vol. 12, p. 99, fig. 1027.

Theridhnn cruciatvm Giebel, 1869, Zeitschr. Ges. Naturwiss., vol. 34, p. 303

Steatoda hipunctata, Thorell, 1856, Nova Acta reg. Soc. sci. Upsaliensis
ser. 3, vol. 2, p. 140. Keyserling, 1884, Die Spinnen Amerikas, vol. 2
pt. 1, p. 116, pi. 6, fig. 76,9,5. Petrunkevitch, 1911, Bull. Amer. Mus
Nat. Hist., vol. 29, p. 187. Wiehle, 1937, in Dahl, Die Tierwelt Deutsch
lauds, pt. 33, p. 193, figs. 200-204,9,(5. Kurata, 1939, Canadian Ent.
vol. 53, p. 81. Roewer, 1942, Katalog der Araneae, vol. 1, p. 412
Gertsch, 1946, in Procter, Biological Survey of the Mount Desert
Eegion, pt. 7; Gertsch, 1949, American spiders, p. 258. Locket and
Millidge, 1953, British spiders, vol. 2, p. 56, figs. 38,9.5. Hackman,
1954, Acta Zool. Fennica, vol. 79, p. 4.
Types. The type locality is presumably Sweden.
Diagnosis. This species may have the legs very slightly shorter

Map 16. American distribution of Steatoda bipunctata (Linnaeus).

than in other species of Steatoda. It is readily distinguished by
comparison of the genitalia (Figs. 89, 155, 156). The total length
of females is 4.8-7.3 mm., of males, 4.4-6.0 mm.

Natural History. This species may have been introduced in
recent times. It seems to have survived mainly along the coast
or the shores of Lake Ontario. C. Dondale has collected speci-
mens on a barn.

Distribution. Europe, Siberia, Kamchatka (Wiehle, 1937).
Introduced in French Guiana and Venezuela (Keyserling, 1884).

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 415

Probably introduced on the coast of Maine to Newfoundland,
north shore of Lake Ontario (Map 16).

Records. Newfoundland: (liackman, 1954) ; Nicholsville. iVoyc
Scotia: Weymouth; Grand Pre; Baddeck; Barrington; Lequille.
New Brunswick: Grand Manan Isl. Quebec: Bonaventura Isl.
Ontario: Toronto (W. J. Gertsch) ; Lakeport. Maine: Cumber-
land Co. : Sebago Lake ; Hancock Co. : Mt. Desert Isl. ; Lincoln
Co.: Jefferson; York Co.: Saeo. New Hampshire: Coos Co.:
Gorham, 1946, (E. L. Bell).

Steatoda grandis Banks

Figures 92-97, 119-123; Map 17.

Steatoda grandis Banks, 1901, Proe. Acad. Nat. Sci. Philadelphia, p. 578;

1910, Bull. U.S. Natl. Mus., no. 72, p. 21, Petiunkevitch, 1911, Bull.

Anier. Mus. Nat. Hist., vol. 2l», p. 18S. Eoewer, 1942, Katalog der

Araneae, vol. 1, p. 413.
Steatoda zionis Chamberlin and Tvie, 1935, Bull. TTuir. Utah, bid. ser. vol.

2, no. 8, p. 12, figs. 3G, 42.9,c5. Roewer, 1942, Katalog der Araneae,

vol. 1, p. 414. NEW SYNONYMY.
Steatoda mcrala I. Fox 1940, Proc. Biol. Soc. Washington, vol. 53, p. 41,

fig. 3, 9 . NEW SYNONYMY.
Types. Female type of S. grandis from Albuquerque, New
Mexico, in the Museum of Comparative Zoology. Male holotype,
female allotype of 8. zionis from Zion National Park, Utah, in
the American IMuseum of Natural History. Female type of 8.
merula from ]\It. Lemmon, Arizona, in the United States Na-
tional Museum (no. 1368).

Diagnosis and Variation. The epigyna of this species are ex-
tremely variable, particularly in the southern portion of its
range (Figs. 93-97). The epigynum of the type of 8. grandis
resembles Figure 93. Only in the narrower septum and slightly
deeper depression on each side of the epigynum does this species
differ from S. americana; furthermore the depression of the
epigynum is frequently filled with a material difficult or impos-
sible to remove. The epigyna of the types of both 8. grandis and
*S'. merula are partly filled in this way. The palpus, in contrast
to the female genitalia, shows little variation (Figs. 119, 120,
123). Only two males from central Colorado had the tip of the
radix slightly different (Figs. 121, 122). Total length of females
4.9-9.0 mm. ; that of males 4.3-7.4 mm.

416

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Natural History. In Colorado this species has been collected
under stones, rocks and in cliffs from lower elevations (7000-
8000 ft.) than S. hespera.

Dist7-ihution. Western United States (Map 17).

Records. South Dakota: Custer Co.: Blue Bell, 4900 ft., $ .
Pennington Co. : Horsethief Lk., 9 ; S. of Rapid City, 9 . Wyom-
ing: Devil's Tower Natl. Mon., Aug. 5, 1952 (B. Malkin), 9 .

S. atascadera "
S grandis •

S palomaro a

1^4

Map 17. Distribution of Steatoda grandis Banks, S. palomara Chamber-
]in and Ivie and S. atascadera Chamberlin and Ivie.

Colorado: Archuleta Co. : Piedra, 9 . Boulder, 9 . Denver, 9 .
Douglas Co. : Parker, 9 . El Paso Co. : Cascade, 9 . Gunnison
Co. : Taylor River, 9 . La Plata Co. : Durango, 9 . Larimer Co. :
Estes Park, 7800 ft., S ; Fort Collins, 9 . Mineral Co. : West
Fork of Wolf Creek, 7800 ft., 9 . Rio Grande Co. : Monte
Vista, 9 . Routt Co. : Steamboat Springs, 9 . Teller Co. : Pikes
Peak, 10,000 ft., 9 . New Mexico: Colfax Co.: Raton, <5 . Ber-
nalillo Co. : Sandia Mts., 9 . Catron Co. : 25 mi. N. of Alma, 9 .

LEVI : SPIDER GENERA CRLTSTULINA AND STEATODA 417

Lincoln Co., 9 . Santa Fe Co. : 10 mi. S. of Santa Fe ; Little
Tesugue Canyon, S . Valencia Co. : Bluewater. Utah : Carbon
Co. : Helper, $ ; Price, 9 , £ . Iron Co. : Parowan, $ S . Emery
Co. : Ferron, 9 , $ . Garfield Co. : Esealante, S ; Red Canyon,

9 ,S . Millard Co. : Scipio, 9 . San Juan Co. : Monticello, 9 .
Sevier Co. : Fish Lk., 9 ; Richfield, 9 ,$ . Uintah Co. : Vernal,

9 ; Watson, 9 ; White Riv. nr. Evacuation Cr., 5000 ft., 9 .
Utah Co. : Levan, 9 . Wayne Co. : Fruita, 9 ; Notom, 9 , $ . Ari-
zona: Apache Co. : Wide Ruin, $ , ,J ; 17 mi. NE. of Whiteriver,

9 , S ; Hannagan, ? . Cochise Co.: Chiricahua Mts. (many col-
lections) 9 , S ; Carr Canyon, Huachuca Mts., $ . Coconino Co.:
Oak Cr. Canyon, 9 . Gila Co. : 7 mi. N. of Payson, 2 ; Middle
Pioneer Cr., Pinal Mts., 9 , S . Graham Co. : Graham Mt. (many
collections) 9 , S . Pima Co. : Santa Catalina Mts. (many col-
lections) 9 ,S . Santa Cruz Co. : Santa Rita Mts. (many collec-
tions), 9 , $ . Yavapai Co. : Prescott, 9 . Oregon: Wheeler Co. :
Richmond, Fall 1948 (Donnely),? ; 3 Aug. 1951 (V. Roth),

9,3.

Steatoda mexicana, new name

Figures 98-103, 124-128 ; Map 18.

Stearodea amerioana F.O.P. Cambridge, 1902, Biologia Centrali-Americana,
Araneidea, vol. 2, p. 377, pi. 35, fig. 12,$. Not Steatoda americana
(Emerton).
Steatoda americana, Petruukevitch, 1911, Bull. Amer. Mus. Nat. Hist., vol.
29, p. 187. Eoewer, 1942, Katalog der Araneae, vol. 1, p. 413. Not
Steatoda americana (Emerton). Not Banks, 1909, Proc. Acad. Nat.
Sci., Philadelphia, vol. 61, p. 205.
Steatoda c/randis, Chamberlin and Ivie, 1935, Bull. Univ. Utah, bid. ser.,
vol. 2, no. 8, figs. 37, 40, 41, $,5. Not S. grandis Banks.
Types. Two female syntypes from Omiltemi, Guerrero in the
British Museum (Natural History).

Diagnosis and Variation. The epigyna of this species are very
variable particularly in the northern and eastern portion of its
range (Figs. 99-103). The epigynum of a syntype examined
resembles that from St. George, Utah (Fig. 99) but the scape
is slightly wider. However, it is slightly narrower than that
from Chiricahua Mountains (Fig. 103). Most females can
readily be told from S. grandis by the wider scape of the epigy-

418

BULLETIN : MUSEUM OP COMPARATH^ ZOOLOGY

num. The shape of the radix of the palpus is variable. Some
almost resemble that of S. grandis, in others the radix is long
and stretched. The shape of the base of the embolus (Fig. 128)
readily distinguishes this species from *S'. hespera. Total length
of females 5.1-9.0 mm. ; of males 4.8-5.7 mm.

Distribution. Idaho to Texas, Arizona south to Guerrero
(Map 18).

Records. Texas: Big Bend Natl. Pk., Chisos Mts., $ . Walker
Co.: Huntsville, 2 Oct. 1950 (M. A. Cazier), 9 . Colorado: Lari-
mer Co.: Eed Feather Lakes, June 14, 1933 (A. B. Klots) 5.
New Mexico: Otero Co. : Camp Mary White, 9 , $ . Idaho: Bear
Lake Co. : Bloomington, 9,5. Utah : Bryce Canyon Natl. Pk., $ ;
Garfield Co. ; 15 mi. N. of Boulder, ? ; Henry Mts., 9 ; 10 mi. SE.

Map 18. Distribution of Steatoda mexicana, new name.

of Panguiteh, 9 . Salt Lake City, 9 . Sevier Co. : Fish Lk., 9 .
Utah Co. : Lehi, 9 ; west shore of Utah Lk., 9 , ^ . Washington
Co. : St. George, 9 ; Beaver Dam Wash., 9 . Arizona: Cochise
Co.: Rustler's Camp. Chiricahua Mts., 9 , c? . Coconino Co.:
Flagstaff, 9 . Grand Canyon Natl. Pk., 9 . Graham Co. : Gra-
liara Mt., Pinecrest, 9 , cJ . Santa Cruz Co. : 4 mi. SE. of Ruliy, 9 .

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 419

Yavapai Co.: Prescott, 9 . Chihuahua: SW. of Santa Barbara.
8400 ft., 9. Durango: " Providencia, " 7700 ft., $ .

Steatoda palomara Chamberlin and Ivie
Figures 104, 105, 129-133; Map 17.

steatoda palomara Chamberlin and Ivie, 1935, Bull. Univ. Utah, biol. ser.,
vol. 2, no. 8, p. 13, fig. 43, $ . Roewer, 1942, Katalog der Araneae, vol.
1, p. 414.

Type. Female type from Mt. Palomar, California in the
American Museum of Natural History.

Diagnosis. The epigynum (Fig. 105) and the tip of the radix
of the palpus (Figs. 130-132) distinguish this species. Total
length of females 5.0-6.8 mm. ; males, 5.3 mm.

Distribution. Known only from San Diego County, California
(Map 17).

Records. California: San Diego Co. : Mt. Palomar, July 25,
1931 (W. Ivie), 9 ; July 13, 1953 (W. J. and J. W. Gertsch), 9 ;
3000-5000 ft., June 30, 1956 (W. J. Gertsch and V. Roth), 9 , S ;
Descanso, June 25, 1947 (W. M. Pearce).

Steatoda atascadera Chamberlin and Ivie
Figures 106-109, 134-141 ; Map 17.

Steatoda atascadera Chamberlin and Ivie, 1942, Bull. Univ. Utah, biol. ser.,
vol. 7, no. 1, p. 39, figs. 83, 84, 9, o.

Types. Male holotype, female allotype from Atascadero Lake,
San Luis Obispo County, California, from under concrete
bridge at south end of lake, in the American Museum of Natural
History.

Diagnosis and Variation. The epigyna (Figs. 107-109) and
palpi (Figs. 134-141) of this species are extremely variable.
Hardly two look alike. The tip of the radix of the male palp
as well as the width of the anterior raised lobe of the epigynum
separate this species from 8. palomara and 8. hespera. Total
length of females 4.8-6.0 mm., of males, 3.8-4.4 mm.

Distribution. Coast range of California from Santa Cruz to
Riverside County (Map 17).

Records. California: Los Angeles Co.: nr. Claremont; (V.
Roth. W. J. Gertsch), 9. San Gabriel Mts. (many records).

420 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

9,$ ; Santa Monica Mts. (R. X. Schick), 9. Glendale (E. I.
Schlinger), 9 . Monterey Co.: Hastings Nat. Hist. Reserve (J.
Linsdale ) , 5 . Orange Co.: Silverado Canyon, Santa Ana Mts.
(R. X. Schick) 9 . Riverside Co. : San Jacinto Mts. (R. X.
Schick), 9 . Santa Cruz Co. : Big Trees Park (A. F. Archer), $ :
Brookdale (W. H. Irvin),5 . Yolo Co.: Davis (E. I. Schlinger).
9.

Steatoda hespeba Chamberlin and Ivie
Figures 90, 91, 110-115, 142-147; Map 19.

Steatoda hespera Chamberlin and Ivie, 1933, Bull. Univ. Utah, biol. ser.,
vol. 2, no. 2, p. 9, figs. 4-6, 9,5. Gertseh, 1939, Amer. Mus. Novitates.
no. 1032, p. 4. Chamberlin and Ivie, 1941, Bull. Univ. Utah, biol, ser.,
vol. 6, no. 3, p. 12. Roewer, 1942, Katalog der Araneae vol. 1, p. 414.
Levi and Levi, 1951, Zoologica, vol. 36, p. 220; 1955, Canadian Field
Nat., vol. 69, p. 69. Lowrie and Gertseh, 1955, Amer. Mus. Novitates,
no. 1736, p. 7.
Types. Syntypes from Clear Creek, Raft River Mountains,
Utah, in the American Museum of Natural History.

Diagnosis and Variation. The proportions of the anterior
raised lobe as well as the even width of the depressed area of the
epigynum (Figs. 111-115) distinguish this species from S. hore-
alis and S. atascadera. The radix of the palpus lacks the median
lobe typical of S. horealis and lacks the lobe below the tip typi-
cal of 8. atascadera (Figs. 143, 146). Specimens from the
western part of the range seem slightly larger in size than those
of the east. Total length of females 4.2-7.5 mm. ; of males,
3.6-5.4 mm.

Natural History. In Colorado this species has been collected
under bark of dead conifers and in cabins at elevations between
8500 ft. and timberline.

Distribution. Western United States (Map 19).
Records. British Columhia: Salmon Arm; Vernon. Montana:
Helena ; Flathead Co. ; Gallatin Co. ; Glacier Natl. Pk. ; Ravalli
Co. Wyomi7ig : Grand Teton Natl. Pk. ; Sublette Co. ; Teton Co. ;
Yellowstone Natl. Pk. Colorado: Gunnison Co.; La Plata Co.;
Pitkin Co. ; Routt Co. Idaho: Adams Co. ; Bear Lake Co. : Frank-
lin Co.; Idaho Co.; Payette Co.; Teton Co.; Twin Falls Co.;

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA

421

Valley Co. Utah: Box Elder Co.; Cache Co.; Garfield Co.;
Millard Co. ; Rich Co. ; Salt Lake Co. ; Sevier Co. ; Summit Co. ;
Utah Co.; Wasatch Co. Nevada: Clark Co.; Reno. Washington:
Ferry Co.; Mt. Ranier Natl. Pk. ; Spokane Co.; Walla Walla;
Whitman Co.; Yakima. Oregon: Baker Co.; Crater Lk. Natl.
Pk. ; Deschutes Co. ; Grant Co. ; Jackson Co. ; Klamath Co. ; Lake
Co. ; Lane Co. ; Morrow Co. ; Three Sisters Primitive Area ;

-T-'^

-^^.J

U

Map. 19. Distribution of Steatoda hespera Chamberlin and Ivie and
6'. borealis (Hentz).

Wheeler Co. California: Calaveras Co.; Contra Costa Co.;
Eldorado Co. ; Fresno Co. ; Kings Canyon Natl. Pk. ; Los Angeles
Co. ; Madera Co. ; Mariposa Co. ; Mendocino Co. ; Modoc Co. ;
Mono Co. ; Monterey Co. ; Nevada Co. ; Placer Co. ; Sacramento
Co. ; Santa Clara Co. ; Santa Cruz Co. ; Sequoia Natl. Pk. ; Shasta
Co. ; Sierra Co. ; Siskiyou Co. ; Tulare Co. ; Yolo Co. ; Yosemite
Natl. Pk.; Yuba Co.

422 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Steatoda borealis (Hentz)
Figures 116-118, 148-154; Map 19.

Theridion boreale Hentz, 1850, Jour. Boston Soc. Nat. Hist., vol. 6, p. 274,
pi. 9, fig. 4,5 ; 1875, Spiders of the United States, p. 145, pi. 16, fig.

4,5 pl.21, fig. 13,5.
Steatoda horcalis, Emerton, 18S2, Trans. Connecticut Acad. Sci., vol. 6,
p. 19, pi. 4, fig. 1,$,5. Keyserling, 1884. Die Spinnen Amerikas, vol.
2, no. 1, p. 119, pi. 6, fig. 77,?, 5. McCook, 1890, American Spiders,
vol. 2, p. 55. Marx, 1890, Proc. U.S. Natl. Mus., vol. 12, p. 521; 1892,
Proc. Ent. Soc. Washington, vol. 2, p. 156. Fox, W., op. cit., p. 268.
Banks, 1892, Proc. Acad. Nat. Sci. Philadelphia, p. 31. Emerton,
1894, Trans. Connecticut Acad. Sci., vol. 9, p. 406. Baker, 1894, Ent.
News, vol. 5, p. 164. Banks, lS95a, Jour. New York Ent. Soc, vol. 3,
p. 84; 1895b, Ent. News, vol. 6, p. 182; 1899, Proc. Ent. Soc. Wash-
ington, vol. 4, p. 189. Emerton, 1902, The common spiders, p. 119, figs.
276-279, 9,6 . Scheffer, 1905, Trans. Kansas Acad. Sci., vol. 19, p. 192.
Banks, 1907, Ann. Eept. Indiana Dept. Geol. and Nat. Res. no. 31, p.
739. Bryant, 1908, Oceas. Papers Boston Soc. Nat. Hist., no. 9, p. 16.
Banks, 1910, Bull. U.S. Natl. Mus., no. 72, p. 21. Petrunkevitch, 1911,
Bull. Amer. Mus. Nat. Hist., vol. 29, p. 188. Banks, 1911, Proc. Acad.
Nat. Sci. Philadelphia, vol. 63, p. 445. Comstock, 1912, The spider
book, p. 360, fig. 363,9. Emerton, 1913, Appalachia, vol. 12, p. 155.
Barrows, 1918, Ohio Jour. Sci., vol. 18, p. 304. Bishop, 1924, Bull. New
York State Mus. no. 251, p. 21. Barrows, 1925, Ann. Ent. Soc. Amer.,
vol. 18, p. 506. Emerton, 1928, Univ. Toronto Studies, Biol., vol. 32, p.
45. Crosby and Bishop, 1928, Mem. Cornell Univ. Agr. Exp. Sta., no.
101, p. 1040. Emerton, 1930, Publ. Nantucket Maria Mitchell Assoc,
vol. 3, p. 164. Worley, 1931, Studies Univ. Nebraska, vol. 27, p. 28.
Chickering, 1931, Papers Michigan Acad. Sci., vol. 15, p. 351. Elliott,

1932, Proc. Indiana Acad. Sci., vol. 41, p. 424. Chamberlin and Ivie,

1933, Bull. Univ. Utah, biol. ser., vol. 2, no. 2, fig. 7, 9 . Stiles and
Detwiler, 1938, Proc. Iowa Acad. Sci., vol. 45, p. 286. Kurata, 1939,
Canadian Nat., vol. 53, p. 81. Comstock, 1940, The spider book, rev. ed.
p. 375; fig. 363,9. Kurata, 1941, Univ. Toronto Studies, biol. ser., no.
48, p. 109. Truman, 1942, Bull. Univ. Pittsburgh, vol. 38, no. 2, p. 4.
Roewer, 1942, Katalog der Araneae, vol. 1, p. 413. Lowrie, 1942, Bull.
Chicago Acad. Sci., vol. 6, p. 169. Kurata, 1943, Canadian Ent., vol. 57.
p. 10. Muma, 1943, Common spiders of Maryland, p. 65, pi. 4, fig. 2,
pi. 12, fig. 20. Chamberlin and Ivie, 1944, Bull. Univ. Utah, biol. ser.,
vol. 8, no. 5, p. 46. Muma, 1945, Bull. Agr. Exp. Sta. Univ. Maryland,
no. A38, p. 27. Gertsch, 1946, in Procter, Biol. Surv. of the Mount
Desert Reg., pt. 7, p. 519. Rapp, 1946, Bull. Brooklyn Ent. Soc.,- vol.

LEVI : SPIDER GENERA CRUSTULINA AND STEATODA 423

41, p. 4. Chamberlin and Ivie, 1947, Bull. Univ. Utah, biol. ser., vol. 10,

no. 3, p. 27. Lowrie, 1948. Ecology, vol. 29, p. 350. Kaston, 1948, Bull.

Connectifut Geol. Nat. Hist. Surv., no. 70, p. 85, figs. 51-56, 2014, $,,5.

Gertseh, 1949, American spiders, p. 258. Kurata, 1949, Canadian Ent.,

vol. 81, p. 127. Elliott, 1953, Proc. Indiana Acad. Sci., vol. 62, p. 309.

Kaston, 1953, How to know the spiders; p. 162. Levi and Field, 1954,

Amer. Midland Nat., vol. 51, p. 444. Kaston, 1955, Trans. Illinois

Acad. Sci., vol. 47, p. 166.

Type locality. "United States." The type specimens are lost.

Diagnosis. The wide raised median lobe, the swollen posterior

area of the depression of the epigynum (Figs. 117-118), and the

median lobe on the palpal radix (Fig. 148) distinguish this

species from S. hespera. Total length of females: 3.8-7.0 mm. ; of

males, 4.3-5.9 mm.

Natural History. This is the common Steatoda of the eastern
United States, which is found on buildings.

Distribution. East of the Rocky Mountains south to Texas
and North Carolina; north to Alaska (Map 19). There are more
old records from the southern states than recent ones. Shreve-
port, Louisiana (Banks, 1899), Alabama (Hentz, 1875) and
Georgia (Keyserling, 1884), however, this species has only rarely
been collected in these states during the last fifty years. One
record probably an accidental one, is that of a male and female
collected at Lakeside, San Diego Count}', California, by N.
Banks, another from Eugene, Oregon, April 1947 (B. Malkin),
$ , probably introduced.

Records. Alaska: (Chamberlin and Ivie. 1947) ; Fox; Tanana:
Mt. McKiiiley Natl. Pk. Nova Scotia: Coldbrook. Quebec:
Montreal; Matapedia. Ontario: (Emerton, 1928; Kurata, 1939,
1941, 1943) ; 36 mi. N. of North Bay; St. Thomas; Lake Tema-
gami; Newmarket; Haliburton. Manitoba: Riding Mountain.
Northwest Territory: (Kurata. 1949), Outpost Island, Great
Slave Lk. Alberta: Medicine Hat; MacMurray; Seba Beach;
Mt. Sentinel. Maine: (Bishop, 1924) ; Cumberland Co.; Piscata-
quis Co. New Hampshire: Carroll Co.; Cheshire Co.; Coos Co.;
Grafton Co. Vermont: Addison Co. Massachusetts: (Emerton,
1930); Es.sex Co.; Middlesex Co.; Norfolk Co.; Plymouth Co.;
Worcester Co. Rhode Island : Vvoxidence. Connecticut: (Kaston,
1948). New York: (Banks, 1892; Bishop and Crosby, 1928);
Clinton Co. ; Fulton Co. ; Jetferson Co. ; Nassau Co. ; Rockland

424 BULLETIN : MUSEUM OF COMPABATIVE ZOOLOGY

Co.; St. Lawrence Co.; Steuben Co. New Jersey: (Rapp. 1946) ;
Bergen Co. ; Cape May Co. ; Mercer Co. ; Morris Co. Pennsyl-
vania: Bradford Co.; Montgomery Co.; Philadelphia Co.; Pitts-
burgh; Schuykill Co. Delaware: Wilmington. Maryland:
(Muma, 1943, 1945). Virginia: Fairfax Co.; Pulaski Co. Wesi
Virginia: Ohio Co. North Carolina: (Banks, 1911) ; Avery Co.;
Buncombe Co.; Guilford Co.; Swaine Co. Mississippi: Rankin
Co. Ohio: Athens Co.; Columbiana Co.; Cuyahoga Co. Michigan:
(Hentz, 1875; Baker, 1894; Chickering, 1931; Lowrie, 1948);
Chippewa Co.; Ogemaw Co.; Washtenaw Co. Indiana: (Banks,
1907; Lowrie, 1948; Elliott, 1953). Kentucky: Hardin Co. Wis-
consin: (Levi and Field, 1954); Crawford Co.; Grant Co.;
Marathon Co.; Milwaukee Co.; Racine Co. Illinois: (Lowrie,
1948, Kaston, 1955); Champaign Co.; Cook Co.; Winnebago
Co. Minnesota: Clay Co.; Clearwater Co.; Freeborn Co.; Mar-
shall Co.; Mille Lacs Co.; Minneapolis; Saint Louis Co. Iowa:
(Stiles and Detwiler, 1938) ; Ames; Cerro Gordo Co.; Woodbury
Co. Missouri: (Banks, 1895b). North Dakota: Divide Co.; Cass
Co. South Dakota: Custer Co. Nebraska: (Worley, 1931) ;
Saline Co. Kansas: Riley Co. Texas: McLellan Co.: Moody,
Sept. 1941 (C. and M. Goodnight),?. Montana: Cascade Co.;
Custer Co.; Daniel Co.; Prairie Co. Colorado: Denver; El Paso
Co. ; Larimer Co. ; Morgan Co.

Figs. 1-3. Crustulina sticta (O.P. Cambridge). 1. Female genitalia,
ventral view. 2. Epigynum. 3. Abdomen of female, dorsal view.

Figs. 4-6. C. altera Gertseh and Archer. 4. Female genitalia, ventral
view. 5. Epigynum. 6. Colulus and spinnerets.

Fig. 7. C. sticta (O.P. Cambridge), left palpus.

Figs. 8-10. C. altera Gertseh and Archer. 8. Palpus. 9. Palpus expanded,
subventral view. 10. Female, dorsal view.

Abbreviations: C, conductor; E, embolus; M, median apophysis; R,
radix; S, subtegulum ; T, tegulum; Y, cymbium.

17

12 /

Figs. 11-13. Stcatoda nigrofemorata (Keyserlmg). 11. Left palpus. 12.
Female genitalia, ventral view. 13. Epigynum.

Fig. 14. S. quaesifa (O.P. Cambridge), palpus.

Figs. 15-18. S. moesfa (O.P. Cambridge). 15. Abdomen of female, dorsal
view. 16. Female genitalia, ventral view. 17. Epigynum. 18. Palpus.

Figs. 19, 20. S. saltensis, new species. 19. Female genitalia, ventral view.
20. Epigynum.

Figs. 21, 22. (S. autumnalis (Banks). 21. Female genitalia, ventral view.
22. Epigynum.

24

28

• v^ #':' '■>^

jiffia^pgg^^^M^^

29

Figs. 23-27. Steatoda transversa (Banks). 23. Female genitalia, ventral
view. 24. Epigynum. 25. Abdomen of female, dorsal view. 26, 27. left
palpus. 26. Mesal view. 27. Ventral view.

Figs. 28-31. S. quadrimaciilata (O.P. Cambridge). 28. Female genitalia,
ventral view. 29. Epigynum. 30, 31. Palpus. 30. (Tamaulipas). 31.
(Guerrero).

36

Figs. '62, 33. Steatoda fulva ( Keyserling j , left palpus. 32. (California).
33. (Florida).

Figs. 34-36. S. meflialis (Banks), palpus. 3-1. (Texas). 35. (Arizona).
36. (California).

Figs. 37-41. S. piilcher (Keyserling), palpus. 37. Mesal view (Cali-
fornia. 38-41. Ventral view. 38. (California K 39. (Texas). 40. (Coa-
huila). 41. (Morelos).

46

44

■."?r*^-i.'

47

^^

St '-•^f^ », -^ :

Ifl^* > ^

1 5.^''

52 53

Figs. 42-44. Steatoda mcdiaU.s (Banks). 42. Female genitalia, ventral
view. 43, 44. Epigynum.

Figs. 45-47. S. fulva (Keyserling). 45. Female genitalia, ventral view.
46, 47. Epigynum.

Figs. 48-51. S. ixdclicr (Keyserling). 48. Female genitalia, ventral view.
49, 50. Epigynmn. 51. Abdomen of a female, dorsal view.

Fig. 52. S. fulva (Keyserling), abdomen of female, dorsal view.

Figs. 53-55. iS'. mcdialis (Banks). 53. Carapace of female. 54, 55.
Alidoinen of female, dorsal view.

Figs. 56-65. Steatoda alboinaculata (De Geer). 56. Carapace of female.
57. Female genitalia, ventral view. 58. Epigynum. 59-61. Left palpus. 60.
Expanded, suljventral view. 61. Expanded, subeetal view. 62. Colulus and
spinnerets. 63. Left ehelieerae of male, inside view. 64-65. Abdomen of
female. 64. Ventral view (Colorado). 65. Dorsal view (Massachusetts).

Ahhrcriations: C, conductor; E, embolus; M, median apophysis; P,
paracymbial hook; R, radix; T, tegulum.

66

^ 71

67

72

Figs. 66-69. Steatoda americana (Enierton). 6Q. Female genitalia, ventral
view. 67. Epigynuni. 68. Left Palpus. 69. Palpus, expanded, subventral
view.

Figs. 70-73. ti. ticptemmaculalu (Keyserling ). 7U. Abdomen of female,
dorsal view. 71. Female genitalia, ventral view. 72. Epigynuni. 73. Palpus.

Abbreviations: C, t-onductoi-; E, embolus; M, median apophysis; E, radix;
T, tegulutn.

.-i^. ^, -■-— '^S?>^''*^'^^'^^^:ii-"'^- .---■

78

81

85

Fig. 74. Steaioda grossa (C.L. Koi-h), left palpus.

Figs. 75, 7^1. <S'. triangulosa (Walfkeiiaei), i>alpus. 7."). Expanded, ectal
view. 7(5. Ventral view.

Figs. 77-79. S. caNtanea (Clerek). 77. I'alpus. 78. Female genitalia
ventral view. 79. Epigyiuiiii.

Figs. 80-82. jS'. triangulo.sa (Walekenaer). 80. Female genitalia, ventral
view. 81. Epigynum. 82. Abdomen of female, dorsal view.

Figs. 83-85. S. grossa (C.L. Koch). 83. Female genitalia, venti-al view.
84. Epigynum (California). 85. Epigynum (Michoacan).

Abbreviations : C, conductor; E, eniliolus; M, median apophysis; R, radix.

Figs. 86-89. Steatoda bipunctata (Linnaeus). 86. Colulus and spinnerets.
87. Left cheliceia of female, inside view. 8S. Female genitalia, dorsal
view. 89. Epigynum.

Figs. 90, 9L S. Jtespera Cliamberlin and Ivie, female. 90. Abdomen,
ventral view. 91. Dorsal view.

'k.'

■ 94

95

96

97

100

101

Figs. 91! 97. Stratuda graiidis Banks. 92. Female genitalia, dorsal view
(Zion Natl. Pk.). 93-97. Epigynuni. 93. (Zion Natl. Pk.). 94. (Richfield,
Utah). 95. (Graham Mts., Arizona). 96, (Hannagan. Arizona). 97. (Alma,
New Mexico).

Figs. 98-103. S. nic.ricuHd, new name. 98. Female genitalia, dorsal view
(Chiricahiia Mts., Arizona). 99-103. Epigynum. 99. (St. George, Utah).
100. (Big Bend, Texas). 101. (Otero Co.. New Mexico). 102. (Utah I>k..
Utah). 103. (Chiricahua Mts., Arizona).

a.fi

104

106

107

108

109

10

114

Figs. lU-t, lUo. Steatoda palomara Chamberlin and Ivie. 104. Female
genitalia, dorsal view. 105. Epigymun.

Figs. 106-109. S. atascadera Chamberlin and Ivie. 106. Female genitalia,
dorsal view. 107-109. Epigynum (all from San Gabriel Mts.).

Figs. 110-115. S. hespera Chamberlin and Ivie. 110, 112, 114. Female
genitalia, dorsal view. Ill, 113, 115. Epigynum. 110, 111. (Montana).
112, 113. (Colorado). 114, 115. (California).

Figs 110-118. .*^. hnrealis (Hentz). 116. Female genitalia, dorsal view.
117, 118. Epigynum. 116, 117. (Wisconsin). 118. (North Carolina).

Figs. 119-123. Steatuda gramlis Banks, left palpus. 119, 120, 123. (Zioii
Natl. Pk.). 119. Mesal view. 120. Ventral view. 121, 122. Radix, ventral
view. 121. (Denver, Colorado). 122. (Estes Park, Colorado). 123. Ectal
view.

Figs. 124-128. -S'. nu.ricana, new name, palpus. 124, 12;'), 128. (Chirica-
hua Mts., Arizona). 124. Mesal view. 125. Ventral view. 126, 127. Radix,
ventral view. 126. (Utah Lk., Utah), 127. (Blooniington, Idaho). 128.
Mesal view.

Figs. 129-133. .S. palomara Clianilierlin and Ivie, palpus (all from Mt.
Palomar, California). 129. Mesal view. 130. Ventral view. 131, 132. Radix
of two different specimens. 133. Ectal view.

Figs. 134-141. Steatoda ntascadera Chamberliii and Ivie, Left palpus.
134, 135, 136. (San Gabriel Mts., California). 134. Mesal view. 135. Ventral
view. 136. Ec'tal view. 137. Radix, ventral view (Fish Canyon). 13S, 139.
(San Gabriel Mts.). 138. Mesal view. 140, 141. (Santa Cruz Co.). 140.
Mesal view. 141. Radix, ventral view.

Figs. 142-147. S. hcspera Clianiljerlin and Ivie, palpus. 142-144. (Mon-
tana). 145-147. (California). 142, 145. Mesal view. 143, 146. Ventral
view. 144, 147. Eetal view.

Figs. 148-154. Steafoda horcaUs (Heiitz), left palpus. 148, 149, 152.
(Wisconsin). 148. Mesal view. 149. Ventral view. 152. Eetal view. 150,
151. Radix, ventral view. 150. (South Dakota). 151. (Massachusetts).
153, 154. Expanded. 153. Mesal view. 154. Ectal view.

Figs. 155-156. S. hipunctata (Linnaeus), palpus. 155. Mesal view. 15(3.
Ectal view.

Abbreviations: C, conductor; E, embolus; M, median apophysis; P,
paracymbial hook; R, radix; T, tegulum; Y, cymbium.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 117, No. 4

CONTRIBUTION TO A REVISION OF THE EARTHWORM
FAMILY OCNERODRILIDAE. THE GENUS NEMA TOGENIA .

By G. E. Gates

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

September, 1957

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
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Bulletin of the Museum of Comparative Zoology

AT nAEVARD COLLEGE
Vol. 117, No. 4

CONTRIBUTION TO A REVISION OF THE EARTHWORM
FAMILY OCNERODRILIDAE. THE GENUS NEMA TOGENIA .

By G. E. Gates

CAMBl{]])(iE, .MASS., U.S.A.
!■ R i X T E U FOR T HE M L' S E U -M

September, 1957

No.4 — Contribution to a revision of the earthworm family

Ocnerodrilidae.
The genus Nematogenia.

By G. E. Gates
INTRODUCTION

Since 1900, three supposedly endemic ocnerodrilids have been
added to the long list (cf. Gates, 1942a) of North American
species known only from types or original material. Three
exotic species have been recorded (Gates, 1954, pp. 245-246) from
the mainland or a neighboring island.

The casual spoil of other collecting (as so often in the past)
now has made possible, for the first time in nearly six decades,
some advance in our understanding of an important constituent
of the oligochaete fauna of the continent.

The author's thanks are extended to Dr. P. J. Darlington.
Jr., and to Dr. G. E. Pickford, for the opportunity of studying
these interesting earthworms.

Genus NeMATOGENIA Eisen, 1900

Ooiierodrilm {Nematogenia) Eisen 1900, Piot-. California Ac. Sci., (3) 2, \>.

112. (Type species, Pygmaeodrilus lacuum Beddard, 1893.)
Xematogenia, Michaelseu, 1900, Das Tierieicli, 10, p. 376.
Xcmaiogpnia, Stephenson, 1930, The Oligochaeta, p. 862.

The genus has been defined (Stephenson, 1930) essentially
as follows : Dorsal pores present. Male pores, united with aper-
tures of the single pair of prostates, on xvii. Spermathecae,
adiverticulate, one pair, with pores on or behind 8/9. Gizzards,
two, in vi-vii. Calciferous sacs, one pair, in ix. Metandric.
Accordingly, two, three or four characters contraindicate inclu-
sion of the Hispaniolan species, described below. Admission
to all other ocnerodrilid genera is, by definition, also denied.
Erection of at least two new genera is required by previous prac-
tice. Re-examination of the basis of classification of the family,
how^ever, first seems preferable.

Ocnerodrilid genera have been defined (cf. Stephenson, 1930,
pp. 856-863) by the following characters; closeness of pairing

428 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

of setae, presence or absence of dorsal pores, presence or absence
of ectal union of male deferent ducts with ducts of one pair of
prostates, number of spermathecae and location of the sperma-
thecal pores, number and location of ^zzards, number and loca-
tion of ealciferous glands, presence or absence of diverticula
on the sperraatliecal duct, numljer and location of prostates,
andry (number and location of testes). However, little import-
ance seems to have been attached to somatic structure. Calci-
ferous glands and gizzards are not mentioned in the definition
of Pygmaeodrilus, and species of Gordiodrilus may have a pair
of lateral sacs or a single, median and ventral one, as well as
one, two or no gizzards. Several genera have the same gizzard
or calciferous-gland characters. Dorsal pores are present, in the
Afro-American portion of the family (that with extramural eal-
ciferous glands), only in two species. Setal pairing is not
mentioned in definitions of five of the genera recognized in 1930.
Wide setal pairing, in anterior and middle regions of the body,
has been reported from one species (Seychelles), in middle and
posterior regions of the body, from two species (South America).
Obviously, then, division of the family into genera has been
based mainly on the genital organs.

The reproductive system, in all earthworm families, always
has provided most of the characters for distinguishing and de-
fining species. It is now being found that this system is "most
susceptible to evolutionary change when reproduction is par-
thenogenetic" (Gates, 1956a, p. 227) for parthenogenesis permits
more rapid accumulation of mutations. Such "evolutionary
plasticity of the reproductive system necessitates considerable
caution in use of its structures for taxonomic purposes at genus
or higher level" (Gates, 1956b, p. 30).

Uniquely diagnostic somatic characters, in the Oligochaeta as
known today, are very rare. None has been found in the Ocnero-
(Irilidae. All species herein considered do, however, share a pair
of somatic characters which are unknown in that combination
elsewhere in the family. A combination of three genital char-
acters, also found in each of the species herein considered is,
however, shared with other groups that now have subgeneric or
generic status. Obviously, then, since other characters are un-
available, the genus must be defined, at least primarily, by

GATES: THE GENUS NEMATOGENIA 429

somatic structure, as below. For the present, non-diagnostic but
universally descriptive genital characters are retained in the
definition. Latero-oesophageal hearts in x-xi appear to be charac-
teristic in the family and need no mention.

Defmition. Bithecal, pores at or behind 8/9. Biprostatic, pores
on xvii. Male pores in or close to xvii.

Gizzards, two, in vi-vii. Calciferous glands, lateral, one pair,
in ix. Spermathecae adiverticulate.

Distribiviion. Dominican Republic (Hispaniola).

The genus is known to be represented elsewhere only by a
peregrine species formerly thought to be African but now ap-
parently of American origin. This peregrine species extends
the generic range to include the following : Panama, Costa Rica,
Cameroons, Fernando Po, Belgian Congo, Ceylon.

Key to species of Nematogenia

1. a. Body small, less than three mm thick, dorsal pores

present 2

b. Body larger, more than three mm thick, dorsal pores

lacking 3

'1. a. Spermathecal pores on 8/9, male deferent and prostatic
ducts of a side unite in parietes to open by a common

pore laciium

h. Spermathecal pores midway between 8/9 and eq/ix,

male pores discrete and behind prostatic pores panamaensis

o. a. Typhlosole present 4

b. Typhlosole lacking 5

4. a. Holandric alba

b. Metandric montana

5. a. Intestinal origin in xii magna

b. Intestinal origin in xiv dominicana

External characteristics that might provide a more useful
key for preliminary identification are liable to individual varia-
tion. Until more is known about such differences, reliance on
characters less liable to intraspecific variation is preferable.
Deficiencies in our knowledge of species erected in the previous
century complicate key construction and obviate any attempt to
show phylogenetic relationships.

430 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Nematogenia magna sp. nov.

Loiiia Kucilla, Cordillera Central, Dominican Republic, under grass in open
pine woods at 8-10,000 feet, June 193S, 0-0-1. Dr. P. J. Darlington
Jr. (Loma Rucilla is said to be the highest mountain in the West
Indies.)

External characteristics. Length, 190 -|- mm {-{- 70 mm?).
Diameter, 8 mm. Segments, ca. 400 + (+ 129?). Pigmentation,
unrecognizable (alcoholic preservation). Prostomium, prolobous.
Secondary furrows; one well marked and postequatorial on each
of iii-vii, one less distinct and pre-equatorial on each of v-vi, one
pre-equatorial and one postequatorial deep furrow on each of
viii-xiii. The posteriormost secondary annulus on each of ix-xi
has a tertiary furrow. Nephropores and dorsal pores unrecog-
nizable, the latter probably lacking.

Setae deeply retracted on preclitellar and clitellar segments,
often unrecognizable, apparently small and not closely paired.
Shortly behind the clitellum, AB=CD<BC<AA, DDyyoC,
but farther on AB<iCD<BC <:AA. Still more posteriorly
AB ca.=BC^or slightly < or slightly > CD, AA appears to
be consistently > BC throughout though the difference may be
small. In xvii, a setae probably present but h setae lacking.

Clitellum red (preservation artefact?), annular, on xiv-xxviii,
the posteriormost portion of xiii with a slight red coloration.

Sphermathecal pores, one pair, at or slightly median to B, on
8/9. Female pores widety separated, equatorial, at approximate
sites of apertures of a follicles, on xiv. Male and prostatic pores
discrete, the former diagonally slit-like, the others smaller,
circular and anterior, all at or just lateral to B and in a single,
transverse, smooth, glistening area with distinct anterior and
posterior boundaries about at levels of 16/17 and 17/18. Lateral
margins of the area indistinct but in BC and nearer C. The pores
of each side are in a slight depression within an area of special
tumescence that is most obvious in AB.

No other genital markings.

Internal anatomy. Septum 6/7 slightly muscular, 7/8-10/11
thickly muscular, 11/12 strengthened but translucent.

Gizzards large, in vi and vii. Calciferous glands one pair, in
ix, of solid type (no lumen), large, lateromesially flattened,
thickly discoidal, slightly notched ventrally. Duct fairly thick,

GATES : THE GENUS NEMATOGENIA 431

soft, from pointed dorsal apex of each gland to lateral face of
gut shortly in front of 9/10, with rather large aperture into
gut lumen. Longitudinal lamellae on inner wall of gut in front
of gland apertures gradually disappear anteriorly in ix. In-
testinal origin in xii. Typhlosole lacking but at mD on roof
of gut for some distance behind Ivi there is a very low ridge,
round to triangular in cross section. Supra-intestinal gland,
in several segments around civ, an obvious yellowish thickening
of gut roof where the tissues have a honey-comb texture.

Dorsal blood vessel single, disappearing from view within the
pharyngeal glands in iii. Ventral trunk unrecognizable anterior
to vii. Subneural lacking or unrecognizable, extra-oesophageals
not distinguishable. Supra-oesophageal recognizable only in ix
where a large branch on each side passes laterally on dorsal face
of calciferous gland stalk and then down into the gland. Commis-
sures of vi-ix obviously connect dorsal and ventral trunks. Com-
missures, or hearts, of x-xi probably latero-oesophageal but there
is no blood in either of the apparent dorsal bifurcations of each
vessel. A small transversely placed body resting on the dorsal
blood vessel but attached to the anterior face of each septum,
from 50/51 posteriorly, resembles the "lymph" glands present
in similar locations of certain species of the megascolecine genus,
Pheretima Kinberg 1866.

Metandric. A longitudinal membrane on each side of the body
between 10/11 and 11/12 just lateral to the testes and male
funnel looks much like the wall of a testis sac but a roof was
not found (testicular coagulum extending well upwards). Sem-
inal vesicles, one pair, in xii, large, lobed. Prostates ca. 20 mm
long, would reach into xliv at least if straight. A duct is not
clearly marked oif but may be about 3 mm long. The male
deferent duct comes into contact with lateral face of the prostatic
duct in xvii but is continued on slightly, somewhat thickened,
before turning mesially to pass into the parietes just behind the
prostatic duct.

Spermathecae, without diverticula or seminal chambers, ca.
6 mm long, flattened against the parietes back to 9/10 and with
a slightly wider portion vertically flattened against anterior
face of 9/10 to which the spermatheca is bound by connective
tissue. Duct and ampulla are not distinctly demarcated ex-

432 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

ternally. If an ectal portion with thick wall and very small
lumen is to be regarded as the duct, it then is much shorter
than the ampulla. A terminal ental portion of the spermatheca
is distended by a white coagulum.

Remarks. A tail portion had been recently broken o&, pre-
sumably during collection. The hind end that was in the same
tube may well be of the same worm though the jagged edges
of the breaks do not exactly fit together. The clitellum is
cracked in several places as often happens when worms are pre-
served with that organ in maximal tumescence. Except for local
macerations of intestinal wall the specimen otherwise is in good
condition.

A supra-intestinal gland of any kind has been unknown
hitherto in any ocnerodrilid. Its anatomy in N. magna needs
characterization from microtome rather than freehand sections
but now appears to be similar to that of the "grid" at end of
the typhlosole in the octochaetine Barogaster Gates 1939. Supra-
intestinal glands have not as yet been looked for in many
earthworm genera. Hitherto the glands usually have been associ-
ated with the posterior end of a median typhlosole. In Nemato-
genia, however, the glands are present only when there is no
typhlosole.

N. magna is distinguishable from all ocnerodrilids, except two
species of Kerriona Michaelsen 1924 and one of Paulistus
Michaelsen 1925, by its size as well as segment number, and from
the species of Kerriona and Paulistus by the presence of two
gizzards (instead of none). A size of 260 x 8 mm, in a family
where all species hitherto known, except those just mentioned,
are 15-60 x 1-3 mm certainly is significant. Recognition of addi-
tional differences from other species of Nemafogcnia awaits more
adequate characterization of those previously erected.

A species, when understood to be an interbreeding population
of organisms with more or less individual if not also geographic
variation, obviously cannot be defined from a single individual.
Nevertheless, some sort of a concise characterization is needed.
In such a precis it is also convenient to list characters, presum-
ably of taxonomic importance, about which information is de-
sired.

GATES : THE GENUS NEMATOGENIA 433

Precis. Spermathecal pores minute, superficial, slightly median
to B, at 8/9. Female pores at sites of missing a setae. Male
pores at or just lateral to B and shortly behind prostatic pores,
both pores of a side in an area of special tumescence within a
median field reaching laterally into BC. Clitelium, annular, on
xiv-xxviii. Setae, not closely paired. (Nephropores? No dorsal
pores.) Prostomium prolobous. (Pigmentation?) Segments, ca.
529. Size, 260 x 8 mm.

Intestinal origin in xii. (No typhlosole.) Supra-intestinal
gland in region around 155th segment. Lymph glands present
posteriorly. Metandric. Spermathecal duct shorter than am-
pulla.

Nematogenia dominicana sp. nov.

San Cristobal, Santo Domingo. From earth of damp forested hillside,
July 19, 1930, 1-0-0. Damp soil in hills to west, July 20, 1930, 0-1-0.
R. Bond per Dr. G. E. Pickford.

External characteristics. Length, ca. 195 mm. Diameter, 5+
mm. Segments, ca. 675 (No. 2). Pigmentation lacking ("gray-
ish pink or pink, paler posteriorly," collector's note). Pro-
stomium, broadly prolobous (No. 1) or indenting i (No. 2) and
then slightly proepilobous. Secondary furrows; one deep and
postsetal on each of vii-xiii, one deep and presetal on each of
viii-xiii, less obvious pre- and postsetal secondary furrows recog-
nizable back to xl. Each pre- and postsetal secondary annulus
of ix-xi has a well marked tertiary furrow\ Nephropores un-
recognizable. Dorsal pores lacking.

Setae first certainly recognizable on ii or iii, very closely
paired, AB ca.=C'Z>, BC much<AA, DD>Y2^, in xxv, BC :
AA : :5 :7 gradually changing to 1 :2 posteriorly, and thence on
constant.

Spermathecal pores, one pair, at B, on 8/9. Female pores un-
recognizable but possibly equatorial and in AB (a present) on
xiv. Male and prostatic pores discrete, the former minute, circu-
lar, the others transversely slit-like and anterior, all about at B
on xvii. Porophores transversely elliptical, flat but markedly
protuberant, with center about at B, reaching into AA and to
17/18 but not to 16/17. A distinctly demarcated, smooth, small.

434 BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY

circular portion bearing the prostatic pore appears to be slightly
retractile and the immediate margin of the pore is raised in a
teat-like protuberance. The male pore is near the posterior
margin of the porophore.

No other genital markings.

Internal anatomy. Septum 5/6 membranous, 6/7 slightly mus-
cular, 7/8-11/12 thickly muscular and funnel-shaped.

Gizzards well developed, in vi-vii. Calciferous glands one
pair, in ix, of solid type (no lumen), rather heart-shaped, with
slight ventral indentation (not disc-shaped). Intestinal origin
in xiv (2). Typhlosole lacking and no slightest rudiment recog-
nizable from xiv-xl but from xli to Ixvi there is a low, band-like
area the wrinkled surface of which contrasts strikingly with the
smoothness of other portions of the gut wall. The band gradually
disappears but at niD a fine ridging still is recognizable and
gradually passes into the supra-intestinal gland, a wide thicken-
ing of gut roof also ridged on veuti-al side. The gland extends
from 180/181 to 194/195 (No. 1, ca. 365 segments) or from
187/188 to 208/209 (No. 2). The gland is slightly constricted
at septal insertions (the juvenile) and has an appearance of
being marked off into segmental glands.

Dorsal blood vessel single, disappearing from view in tissue
masses of the pharyngeal bulb that reach to 5/6, in intestinal
region receiving in each segment two pairs of large vessels from
the gut, one pair near each septum. Supra-oesophageal trunk
recognizable only in ix-xi, apparently ending with hearts of
xi. Extra-oesophageals filled with blood and recognizable only
in viii-ix where they are median to the hearts and free from
the gut. A large vessel passes out from the supra-oesopliageal
in ix, on each side, and shortly bifurcates, one branch passing
along calciferous duct to and into the gland, the other branch
passing down to join the extra-oesophageal. Subneural quite
unrecognizable (probably lacking?). Segmental commissures
connecting dorsal and ventral trunks quite slender in v-vii, some-
what larger in viii-ix (lateral hearts?). Hearts of x-xi appar
ently latero-oesophageal but blood present only in the bifurca-
tions that pass to the supra-oesophageal. Lymph glands present
and apparently as in magna.

GATES: THE GENUS NEMATOGENIA 435

Metandrie. Seminal vesicles, one pair, in xii. Prostates 18-20
mm long, would reach at least into xliv if straight, pink (for-
malin developed artefact?). An ectal white portion, slightly
slenderer, 1-2 mm long, with marked muscular sheen constitutes
the duct. The male deferent duct is thicker behind 16/17, in
contact with lateral face of prostatic duct but continued on as in
other species and is narrowed again within the parietes.

Spermathecae, 4+ nim long, without diverticula or seminal
chambers, but with distinctly demarcated duct and ampulla.
The latter, ca. 1 mm long, elliptical in cross section, ca. twice
as thick as ental portion of duct, filled with a white coagulum.
Duct, circular in cross section, very shortly and closely looped
eetally.

Reproduction. Brilliant iridescence on male funnels and in
spermatheeal ducts eetally (but not in coagulum of spermathecal
ampullae) shows that sperm had been matured and that copula-
tion had been completed, in the larger specimen. Reproduction
presumably is sexual and biparental.

Regeneration. Segment length, at 395/396, abruptly becomes
very much shorter and so continues posteriorly, as in a tail re-
generate.

Remarks. The aelitellate worm had been ruptured in the in-
testinal region. No signs of postsexual regression were recog-
nized in that worm which may have been preserved prior to
development of the clitellum. In that case copulation takes
place, as has been suggested for other species, before the clitel-
lum develops.

Absence of a tj^phlosole and presence of a supra-intestinal
gland indicate relationship to magna from which dominicana
is distinguished as follows : By constant close pairing of setae
throughout the Ijody as well as other setal characters. Presence
of paired male porophores. Intestinal origin in xiv (rather than
xii), an important distinction in a family where the origin so
very frequentl}' is in xii. Greater length of spermathecal duct
relative to that of the ampulla. Further differences may be
recognizable when clitellate specimens become available.

Precis. Spermathecal pores minute, superficial, at B, on 8/9.
Female pores in ABl Male pores about at B, shortly behind
prostatic pores, in paired, transversely elliptical porophores.

436 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

(Clitellum?) Setae very closely paired, BC<AA, DD>y2C.
(Nephropores? No dorsal pores.) Prostomium prolobous. Un-
pigmented (?). Segments, 675. Size, 195 x 5 mm.

Intestinal origin in xiv. (No typhlosole.) Supra-intestinal
gland in ten or more segments behind 180/181. Lymph glands
present posteriorly. Metandric. Spermatheeal duct longer than
ampulla, ectally with short loops.

NeMATOGENIA MONTANA Sp. nOV.

Loma Rucilla, Cordillera Central, Dominican Republic, under grass in open
pine woods at 8-10,000 feet, June 1938, 0-0-2. Dr. P. J. Darlington Jr.

External characteristics. Length, 85 and 108 mm. Diameter,
4 mm (clitellum). Segments, 178 and 144 (±1, a metameric
anomaly). Pigmentation, unrecognizable (alcoholic preserva-
tion). Prostomium, prolobous (2 specimens). Secondary fur-
rows, one presetal and one postsetal each on vii-ix. Nephropores
and dorsal pores unrecognizable, the latter probably lacking.

Setae closelj^ paired anterior to clitellum, more widely paired
behind clitellum, farther posteriorly AB ca.=or < CD <BC<A^,
DD^or slightly <y2C.

Clitellum white, annular, from 13/14 onto anterior portion of
xxi (No. 1) or to 21/22 (No. 2), intersegmental furrows present
(No. 1) or lacking (No. 2), setae probably present.

Spermatheeal pores, one pair, about at B, on anterior margin
of ix. Female pores, probably slightly anteromedian to a setae,
on xiv. Male and prostatic pores unrecognizable but presumably
in or near a small indistinctly demarcated white area in AB, on
xvii.

No other genital markings.

Internal anatomy. Septa 5/6-11/12 muscular to thickly mus-
cular.

Gizzards well developed, in vi and vii. Calciferous glands, one
pair, in ix, of solid type (no lumen), sausage-shaped, opening
through a short stalk from dorsal end into oesophagus anterior
to 9/10. Intestinal origin in xiii (2). Typhlosole begins rather
gradually in region of xvii-xxii, reaching maximum height (0.75
mm) in xxv-xxvi, lamelliform, apparently ending abruptly in
Ixxxvii (No. 1) or ciii (No. 2) though more careful examination

GATES: THE GENUS NEMATOGENIA 437

reveals a rudimentary continuation that gradually diminishes
and is unrecognizable behind xcv (No. 1) or cxv (No. 2). Supra-
intestinal gland lacking.

Vascular system, with latero-oesophageal hearts in x-xi, no
subneural, but with "Ij-mph" glands, and, so far as can be
determined, as in magna.

Metaudrie. Seminal vesicles, one pair, lobed, in xii. Prostates
ca. 14 mm long, would reach at least into region of xxxviii-xl if
straight (duet indistinguishable externally), passing into par-
ietes at B. Male deferent duct just lateral to prostate in xvii,
and only well behind ectal portion of that gland turning mesially
to pass, slightly widened, into parietes about at A and just in
front of level of 17/18.

Spermathecae, without diverticula or seminal chambers. Am-
pullae, on anterior face of 9/10, clearly demarcated from a
slender duct that may be nearly as long as the ampulla or some-
what shorter.

Reproduction. Iridescence on male funnels, in spermathecal
ducts and in coagulum of each spermathecal ampulla is brilliant
and demonstrates maturation of sperm as well as completion of
copulation. Reproduction presumably is sexual and biparental.

Regeneration. The penultimate segment of each worm is large
and setigerous. The anal "segment" is unusually small, scarcely
more than a narrow band around a vertically slit-like anal
aperture. Such an anal band, without pigment, develops in
pigmented species shortly after posterior amputation and is
quite different at that stage from the normal anal region.

Segment cii of No. 1 is only about half as long as ci and
succeeding segments for some distance are of the same length.
The region behind 101/102 would have been suspected of being
a regenerate except for the increase in length of the last few
segments.

Parasites. Cysts are scattered through the coelomic cavities
of a posterior portion of the body in one worm.

Remarks. N. montana is clearly distinguished from magna by
the much smaller size, shorter clitellum, presence of a typhlosole.
and absence of a supra-intestinal gland. Distinctions from domin-
icana are : much smaller number of segments, wider pairing of
setae behind the clitellum, shorter and unlooped spermathecal

438 BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY

duct, intestinal origin in xiii instead of xiv, presence of a typhlo-
sole, and absence of a supra-intestinal gland. The only differences
from the inadequate!}^ characterized pmiamaensis that can be
cited at present are: greater size (rather than 35-75 x 1-2 mm),
larger number of segments (rather than 110-120), absence of
dorsal pores, wider pairing of setae behind the clitellum. Pre-
sumably closest relationships are with panamaensis.

Absence of male funnels, or any rudiments thereof, in seg-
ment X of this and the two preceding species appears to be
indicative of long establishment of the metandry.

Precis. Spermathecal pores minute, superficial, about at B,
on anterior margin of ix. Female pores anteromedian to a (?).
Male pores behind prostatic pores which are also in AB (no
porophores or male field?). Clitellum, annular, on xiv-xxii. Setae
more widely paired behind clitellum, AB=or< CD <BC<A^,
i>Z>=or<yoC. (Nephropores? No dorsal pores.) Prostomium
prolobous. (Pigmentation?) Segments, 144-178. Size, 85-105 x
4 mm.

Intestinal origin in xiii. Typhlosole simply lamelliform. (No
supra-intestinal gland.) Lymph glands present posteriorly. Met-
andric. Spermathecal duct, about as long as ampulla, slender.

Nematogenia alba sp. nov.

Lonia EiK'illu, Cordillera Central, Dominican Kepublic, under grass in
open pine woods at 8-10,000 feet, June 1938, 1-3-2. Dr. P. J. Darling-
ton Jr.

External cliar act eristics. Length, 95-115 mm. Diameter, 4-4-f
mm. Segments, 211, 213, 230. Pigmentation, unrecognizable
(alcoholic preservation). Prostomium, tanylobous, a definite
transverse furrow across the tongue shortly behind anterior
margin of i (2 specimens) produces a condition that can be des-
ignated as combined tany- and epiloby. Secondary furrows, one
presetal and one postsetal on some or all of viii-xi. Nephropores
and dorsal pores unrecognizable, the latter probably lacking.

Setae begin on ii and are closely paired, on xx AB ca.=CD<C
BC much<or=J.J,, DD ca.=or slightly < 1/20.

Clitellum red, annular, on xiii-xx (1), setae present, inter-
segmental furrows obliterated except that a slight trace of
13/14 is still recognizable in ventrum.

GATES: THE GENUS NEMATOGENIA 439

Spermathecal pores, one pair, very small, transverse slits, on
or just lateral to B, definitely though only slightly behind 8/9.
Female pores, minute, diagonal slits, in front of a setae. Male
and prostatic pores discrete, in seminal furrows. One limb of
each furrow is about on A between level of 17/18 (unrecogniz-
able in ventrum) and eq/xvii, the other limb passing almost at a
right angle laterally along e(i/xvii to B. The male pore is at
the posterior end of the groove. The prostatic pore has not been
seen but the prostatic duct has been traced through the body
wall to anterior end of the groove. Each seminal furrow is in
a small region of special tumescence reaching from about mBC
median to A, to eq/xviii but not to level of 16/17. Both tumes-
cences, on the clitellate worm, are within a single transverse
field".

No other genital markings.

Internal anatomy. Septum 6/7 membranous or only very
slightly strengthened, (5/6?), 7/8-11/12 slightly muscular and
translucent.

Gizzards well developed, apparently in vi-vii. Calciferous
glands, one pair, in ix, of solid type (no lumen), ovoidal, with
rather short but thick stalk from dorsal apex passing to lateral
face of oesophagus anterior to 9/10. Oesophagus valvular
through xiii (3). Intestinal origin in xiv (3), the real origin
concealed by symmetrical or asymmetrical herniation of intes-
tine through 13/14 well into xiii. Typhlosole rudimentary in
xv-xix, gradually becoming lamelliform and attaining maximum
height (1 + mm) in xix-xx, gradually losing height posteriorly
and unrecognizable behind 158/159 (juvenile with 230 seg-
ments), 140/141 (211 segments, posterior amputee?), or 143/144
(213 segments, apparently with a very small anal-segment re-
generate) and in this worm ending abruptly rather than gradu-
ally and indefinitel3\ Supra-intestinal gland lacking.

Vascular system with latero-oesophageal liearts in x-xi, no sub-
neural ( lymph glands ? ) ; otherwise, so far as can be determined,
as in magna. Nephridia present from ii, ducts (behind the
clitellum) pass into parietes in AB.

Holandric. Seminal vesicles in ix and xii. Those of ix may
be much larger, extending anteriorly into vii but in a worm
with right posterior vesicle continued into xv, the left anterior

440 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

vesicle is confined to ix. Prostates, 17+ mm long, would reach
into Ix if straight, elliptical in cross section and with slit-shaped
lumen. An ectal portion ca. two mm long, with muscular sheen,
circular in cross section, with narrower lumen, is the duct. Male
deferent duct lateral to the prostate in xvii, posteriorly turning
mesially to pass into parietes at level of 17/18 about at A,
slightly thickened just prior to entrance into parietes.

Spermathecal duct about as long as ampulla from which it is
clearly demarcated by a definite constriction, thicker entally and
there elliptical in cross section, with a slit-shaped lumen, ectal
portion circular in cross section and with much smaller lumen.
Coagulum that fills the ampulla is without iridescence. The
iridescence that characterizes the ental portion of the duct is
recognizable in transverse sections only in a jagged circle ap-
parently at the middle of the wall.

Reproduction. Brilliant iridescence on male funnels and in
ental portions of spermathecal ducts (aclitellate and clitellate
worms) shows that sperm had been matured and that copulation
had taken place. Reproduction presumably is sexual and bi-
l)arental.

Remarks. The clitellum of one worm listed as clitellate actu-
ally is only slightly developed.

Prostates of one worm pass straight forward into viii instead
of posteriorly as usual.

The cuticle was found, several years after the Hispaniolan
worms were dissected, to be loosened from the epidermis in vari-
ous small portions of the body. Removal of loosened patches
permitted, in better conditions, another search for nephropores.
None was found in three species. Possible sites in N. alba may
be as follows : At or slightlj^ above D and near anterior segmental
margins, on iii-ix or x. Posteriorly, on segmental equators (but
with some irregularity?) and just median to D.

Male maturity, here as in dominicana, apparently may be
reached and copulation completed before appearance of the
clitellum. The latter organ, at full development, is indicative
primarily of female maturity. Copulation may even be a pre-
requisite for clitellar development.

N . alba is distinguished from all species hitherto referred to
the genus by its holandry, from all American species by the

GATES : THE GENUS NEMATOGENIA 441

presence of seminal grooves, and, with exception of montana, by
the presence of a typhlosole. Further distinctions from montana
are provided by the intestinal origin in xiv, and the prostomium.

Precis. Spermathecal pores small transverse slits, at or just
lateral to B, slightly behind 8/9. Female pores in front of a.
Male pores in posterior ends of seminal grooves, about at A and
17/18. Male field transverse, seminal grooves along A from
17/18 to eq/xvii and thence to B. Clitellum, annular, on xiii-xx.
Setae, closely paired, AB=CD, BC<or=AA, DD ca=y2C.
(Nephropores? No dorsal pores.) Prostomium tanylobous but
with tongue crossed by a transverse furrow. (Pigmentation?)
Segments, 211-230. Size, 95-115 x 4 mm.

Intestinal origin in xiv. Typhlosole vertically lamelliform. (No
supra-intestinal gland. Lymph glands?) Holandric. Seminal
vesicles in ix and xii. Spermathecal duct about as long as
ampulla, thicker entally.

Nematogenia lacuum Beddard, 1893

Fygmacodrilus lacuum Beddard 1893, Quart. J. Micr. Sci., 3-4, p. 259,
figs. 13-16 (pi. 26). (Type loealitj', supposedly Lagos in West Africa.
Types, if extant, in the British Mus.)
OcnerodrUus lacuum, Beddard, 1895, Monog. Oligoehaeta, p. 515.
Xematopenm lacuum, Michaelsen, 1900, Das Tierreich, 10, p. 376.

This taxon, unfortunately, was made the type species. It is
known only from the original description of "five or six" speci-
mens supposedly from Lagos (AVest Africa) but collected alive
at Kew in England. The anatomy was determined from sec-
tions partly ruined by sand which presumal)ly had not been
cleared out from the gut. No information is available as to
nephropores, female pores, male field, typhlosole, supra-intestinal
glands, major blood vessels, hearts.

The species always has been assumed to be endemic in Africa.

Precis. Spermathecal pores in AB, at 8/9. (Female pores?)
Male pores, apertures of united male deferent and prostatic
ducts, in AB, in oval elevations in xvii. Clitellum, saddle-shaped,
on xiii/2-xxvi/2. Setae closely paired, a&/xvii-xviii lacking.
(Nephropores?) Dorsal pores present, beginning at 6/7 or even
anteriorly. Prostomium prolobous( ?). (Pigmentation?) Seg-
ments, ca. 120. Size, 25 (x ]-f ?) mm.

442 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Intestinal origin in xii. (Typhlosole? Supra-intestinal gland?
Lymph glands? Hearts?) Metandric. Seminal vesicles in xii.
Small bulbils ejaeulatorius present.

Spermathecal duct slender, longer than the ampulla.

Dhtribution. Unknown. (See above.)

Nematogenia panamaensis Eisen, 1900

Ocncrodrilus {Nematogenia) lacuuin var. panamaensis, Eiscn 1900, Proc.

California Ac. Sci., (3) 2, p. 127, figs. 55-65 (pi. 9), 114-116 (pi. 11).

(Type locality, Panama. Types, if extant, probably in U. S. Nat. Mus.)
Nematogenia panamaensis, Miehaelsen, 1900, Das Tierreich, 10, p. 376.
Nematogenia panamaensis, Miehaelsen, 1903, Ark. Zool. 1, p. 163. (Camer-

oons)
Nematogenia panamaensis, Miehaelsen, 1904, Sitzber. PJohm. Ges. Prag,

1903, (40), p. 16. (Ceylon)
Nematogenia josepliina Cognetti, 1904, Boll. Mus. Torino, 19, (478), p. 3.

(Type locality, San Jose, Costa Kica. Types in Torino Mus.?)
Nematogenia josepliina, Cognetti, 1906, Mem. Ac. Sci. Torino, 56, p. 55.
Nematogonia (sic) panamaensis, Cognetti, 1908, Ann. Mus. Genova, (3), 4,

p. 33. (Fernando Po)
Xcnuitogenia panamaensis, Miehaelsen, 1915, Ergeb. 2 Deutschen Zentral-

Afrika-Exp. 1910-11, 1, (Zool. 1), p. 222. (Belgian Congo)
Xematogcnia panamaensis, Stephenson, 1923, Oligochaeta, in Fauna Britisli

India, p. 483.
Prostomium, female pores, typlilosole, lymph and supra-in-
testinal glands, hearts and other portions of the vascular system.
in the original description, were not mentioned. Nor have most
of those deticiencies as yet been rectified. A blood vessel in seg-
ment xii is indeed labelled heart (Eisen, 1900, fig. 56h, pi. 9)
but none was shown in xi where a pair certainly is present.
Hearts in xii, in the Ocnerodriliclae, would be so unusual that
abnormality or a mistake in drawing must, in absence of sub-
sequent confirmation, be assumed.

Male pores are discrete. Hence, a statement to the contrary
(and applicable only to lacuum) in the generic definition always
has been unwarranted.

Sperm were seen by Eisen in the spermathecal ampulla. Re-
production, since sperm are matured and exchanged in copula-
tion, can be assumed to be sexual and biparental.

N. josepliina is distinguished from panamaensis, according to
its author (Cognetti, 1906, p. 57) by location of the first dorsal

GATES : THE GENUS NEMATOGENIA 443

pore (at 8/9 instead of 10/11), length of prostates, and absence
of hearts in xii. The first two characters, in better known
species, are subject to considerable individual variation and the
third difference probably is not real. If spermathecal pores were
correctly located by Cognetti, at 8/9, there would be reason
for believing some distinct status eventually might be justified.

iV. panamaensis is peregrine, in the sense usually intended bj^
that word in oligochaetological literature, the overseas discon-
tinuities in its distribution due to transportation, presumably
by man and since the days of Henry the Navigator. The discrete
male pores (if valid evidence of relationship) then require, in-
stead of an African origin as previously maintained, one in
America.

Precis. Spermathecal pores in AB, midway between 8/9 and
eq/ix. (Female pores anterior to a?) Male pores behind pro-
static pores, in a single pair of porophores. Clitellum (shape?),
in xiii, xiii/u, xiv-xxi, xxii/n. Setae, closely paired, a,&/xvii
lacking. Nephropores at D. Dorsal pores beginning at or behind
8/9. (Prostomium? Pigmentation?) Segments 110-120. Size,
37-55 X 1-2 mm.

Intestinal origin in xiii. (Typhlosole? Supra-intestinal and
lymph glands?) Metandric. Spermathecal duct slender, shorter
than ampulla.

Distrihution. Panama, Costa Rica (San Jose), Cameroons
(Bange, Bibundi, Buea, Victoria, Musake), Fernando Po, Bel-
gian Congo (Kongo da Lemba and St. Joseph on the Luala
River), Ceylon (Peradeniya).

'1?

. fJ

Key to genera of the Oc7t,erodrilidae^

1. a. Extramural calciferous glands lacking

h. Extramural calciferous glands present .

•J. (1. One gizzai d, in vii Malabaria, Thatonia.

b. Two gizzards, in vi-vii Deccania.

.">. a. Calciferous sacs paired, lateral 4

h. Califerous sacs unpaired, median and ven-
tral 8

* Malabarin, Thutonin. Deccania arc known only from India and Burma
Maheina onlv from the Seychelles (to which it may have Ix-en transported
recently), ciirgimm only from India (to which it may have l)een introduced
recently). Gordiodriliiii (as of Michaelsen & Stephenson) is African. Other genera
are Arnerican or American and African.

444 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

4. a. Calcif erous sacs in ix and x 5

b. Calcif erous sacs in ix only 6

5. a. One gizzard, in vi Maheina.

b. No gizzard Paulistus, Quechuona.

(i. a. One or more gizzards 7

b. No gizzards Oonerodrilus, including

Liodrilus, Ka-plodrilus,
Ilyogenia, as well as
Pygmaeodrilus, Eu-
Icerria (part), Bia-
phorodrilus, Kerriona.

7. a. One gizzard, in vii Euherria (as it must be

restricted).
b. Two gizzards, in vi-vii Nematogenia.

8. n. Calcif erous sacs in ix-x, (gizzard in vii) . . . Curgiona.
b. Calcif erous sacs in ix 9

9. a. One or more gizzards 10

b. No gizzard Gordiodrilus (part),

as it must be
restricted.

10. a. One gizzard, in viii Gordiodrilus, a

"robustus" group of
species.

b. Two gizzards, in vii-viii Gordiodrilus, a

' ' nannodrihis ' ' group
of species.

DISCUSSION

A taxonomy based primarily on genital structure culminated,
in the Oenerodrilidae, in the congeries known as Gordiodrilus
(cf. Gates, 1942b, pp. 75-85). That taxonomie waste-basket
which defeated Steplienson, "I can do nothing with this hetero-
geneous group" (1930, p. 863), probably could have been
avoided (CTates, 1942b, p. 76) if somatic structure had been
taken into consideration, even as little as previously. Collec-
tions that would justify a real attempt at revision of Gordio-
drilus, unfortunately, are no more available now than in 1942.

Errors of typography, observation and interpretation, so
frequent in the literature on Gordiodrilus, may well be much less
common in the literature on other ocnerodrilid genera. Neverthe-
less, more adequate characterization of somatic systems still is

GATES: THE GENUS NEMATOGENIA 445

prerequisite to a revision of any portion of the family. Also
needed is information as to individual and geographical varia-
tion about which next to nothing is now known.

Some of the taxonomic changes that will be made in th(?
future can be deduced from a key constructed entirely on
characters supplied by just one somatic system. That is the
digestive system. The organs involved are gizzards and calci-
ferous glands, with the glands having the greater importance.

Genera that key out together are presently distinguishable
from each other only by means of characters furnished by the
system usually most liable to rapid evolutionary change. The
genital characters by which those genera are defined and dis-
tinguished, elsewhere in the family and/or in other families, do
not, and sometimes cannot, have that taxonomic value.

Relationships suggested by somatic systems are of course
often different from those that were deduced in the past from
reproductive structure. Thus DiapJiorodrilus now seems to be
closer to Ocnerodrilus than to Gordiodrilus with which it w^as
mistakenly united. Kerriona may be closer to Ocnerodrilus than
to its supposed ancestor Kerria.

REFEEENCES

Gates, G. E.

1942a. Check list and bibliography of North American earthworms.

Am. Midland Nat., 27: 86-108.
1942b. Notes on various peregrine earthworms. Bull. Mus. Comp. Zool.

Harvard, 89: 63-144.
1954. Exotic earthworms of the United States. Bull. Mus. Comp.

Zool. Harvard, 111: 219-258.
1956a. Reproductive organ polymorphism in earthworms of the oriental

megaseoleoine genus Phcretima Kinberg 1867. Evolution, 10:

213-227.
1956b. Notes on American eavthworms of the family Lumbricidae.

III-VIT. Bull. Mus. Comp. Zool. Harvard, 115: 1-46.

Stephenson, J.

1930. The Oligochaeta. Oxford.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE
Vol. 117, No. 5

TRIASSIC REPTILE FOOTPRINT FAUNULES
FROM MILFORD, NEW JERSEY

By Donald Baird

With Four Plates

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

November, 1957

Publications Issued by or in Connection

WITH THE

MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE

Bulletin (octavo) 1863 — The current volume is Vol. 117.

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Vol. 55.

Johnsonia (quarto) 1941 — ^A publication of the Department of
Molliisks. Vol. 3, no. 35 is current.

Occasional Papers of the Department of Mollusks (octavo)
1945 — Vol. 2, no. 21 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-
bers which may be purchased separately. Prices and lists may be
obtained on application to the Director of the Museum of Comparative
Zoology, Cambridge 38, Massachusetts.

Of the Peters ' ' Check List of Birds of the World, ' ' volumes 1-3 are
out of print ; volumes 4 and 6 may be obtained from the Harvard Uni-
versity Press ; volumes 5 and 7 are sold by the Museum, and future
volumes will be published under Museum auspices.

Bulletin of the Museum of Comparative Zoology

AT HARVARD COLLEGE

Vol. 117, No. 5

TRIASSIC REPTILE FOOTPRINT FAUNULES
FROM MILFORD, NEW JERSEY

By Donald Baird

With Four Plates

CAMBRIDGE, MASS., U.S.A.

printed for the museum
November, 1957

No. 5 — Triassic Reptile Footprint Faunules from Milford, New

Jersey

By Donald Baied

CONTENTS

Page

FOREWORD 449

GEOLOGICAL OCCURRENCE 451

SYSTEMATIC DESCRIPTIONS 452

THEROPODA

Grallator sulcatus Baird, n. sp 453

Anchisauripus milfordetisis (Bock) 462

Anchisauriprts parallelus (Hitchcock) 464

Genus incertum 465

Taxonomy of theropod footprints 467

PSEUDOSUCHIA

Chirotherium parvum (G. H. Hitchcock) 473

Chirotherium ey.ermani Baird, n. sp. 479

PHYTOSAURIA

Apatopus lineatus (Bock), n. gen. 486

LACERTOIDEA INCERTAE SEDIS

Rhynchosauroides hyperiates Baird, n. sp 494

Rhynchosauroides sp. 497

CORRELATIVE AGE OF THE MILFORD FAUNULES 501

REPTILES OF THE BRUNSWICK FORMATION 503

APPENDIX I : The nature of Otouphepm 507

APPENDIX II : Brontozoum 509

SUMMARY 511

REFERENCES 514

FOREWORD

In a recent paper Wilhelm Bock (1952) has described a num-
ber of important faunules of fossil footprints from various
horizons and localities in the Upper Triassic (Middle Keuper)
of the Newark Basin. One of these faunules, found in a gray
sandstone layer of the upper Brunswick formation near Milford,
New Jersey, had been discussed in the literature as early as
1886 ; but until 1952 only its dominant species, Chirotherium
parvum, had been adequately described. In the same paper

450 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Bock described a new species of pseudosuchian footprints, Chiro-
therium lulli, found in a reddish-brown siltstone bed at the same
locality. More recently a slab bearing the counterpart trackway
of C. lulli and two forms of dinosaur tracks has come to light
and, with Bock's generous permission, has been described
(Baird, 1954).

Most of the Milford footprints discussed by Bock had been
collected by John Eyerman in 1885 and deposited in the geologi-
cal museum of Lafayette College at Easton, Pennsylvania (here-
after abbreviated LC). One slab, privately owned, had been
acquired by Bock for the Academy of Natural Sciences of Phila-
delphia (ANS). "While Bock's paper was in press, Eyerman 's
second and much larger collection from the same quarries came
to light in the Museum of Comparative Zoology (MCZ), and
received independent preliminary study. This material, though
described in part by Eyerman in 1889, had been overlooked by
subsequent authors. A search of the American Museum of Natu-
ral History collections (AMNH) has produced eight additional
footprint-bearing sial)s collected at Milford by Eyerman in 1886.

Besides increasing the Chirotherium parvum faunule by sev-
eral species and permitting morphological re-interpretation of
those already described, the additional material includes a new
species of dinosaur footprint which represents a third Milford
faunal horizon. As this material also explains certain 19th-
century misidentifications which have distorted the faunal pic-
ture, and the geological correlations which have been based on
it, I have undertaken a comprehensive re-examination of the
Milford reptile footprints and their zoological and stratigraphic
significance.

I am particularly indebted to Wilhelm Bock for photographs
and for permission to mold the type of Chirotherium copei, to
J. L. Dyson for the loan of the Lafayette College types, and to
E. H. Colbert for the loan of American Museum specimens and
for generously making available his unpublished data on Coelo-
physis. Special thanks for the privilege of studying the collec-
tions under their care are due to G. W. Bain, A. E. Wood, and
R. E. Schortmann at Amherst, R. G. Chaffee at Dartmouth, E.
H. Colbert and Mrs. R. H. Nichols at the American Museum,
D. H. Dunkle at the U. S. National Museum, J. T. Gregory at
Yale, J. L. Kay at the Carnegie Museum, and C. W. Waldron,

TRIASSIC REPTILE FOOTPRINTS 451

Jr. at the Museum of Science, Boston. For their helpful and
stimulating susrgfestions 1 am pleasantly indebted to C. L. Camp,

E. H. Colbert, the late R. S. Lull, the late J. H. McGregor,

F. E. Peabody, and D. M. S. Watson. The continued support
and encouragement of Dr. Alfred S. Romer, as well as his
critical reading of the manuscript, are most gratefully acknowl-
edged.

GEOLOGICAL OCCURRENCE

Two distinct footprint horizons bearing mutually exclusive
faunules occur in the gray sandstones of the Milford quarries.
Eyerman nowhere indicates the relative stratigraphic positions
of these horizons, but they are probably not very widely sep-
arated : both are referred to the upper Brunswick formation,
about 5,100 meters above the Triassic base. For clarity they are
here arbitrarily designated Levels A and B. Although this can-
not be stated as certain, Eyerman 's discussion of the MCZ
material (1889, p. 32) implies that the Smith Clark quarry was
the source of the Level B or Chirotherium parvum faunule,
while another quarry about one-half mile to the east furnished
all the material from Level A.

At Level A, the footprints were impressed in gray shale and
are preserved as natural casts on the under surface of a gray
sandstone 4 to 6 cm. thick. The upper surface of this sandstone
is irregularly excavated into shallow pockets in which adhere
pieces of the overlying gray shale. All the footprints are
referred to a single species, Grallator sulcatus. Most are deeply
impressed, between 1 and 2 cm. below surface level at their deep-
est points; many are obscured by slippage, overlapping, or
slumping of the extruded clay. None shows impressions of the
plantar scales and very few show fringe-scale striations. Among
the tracks on MCZ 216 is a striated drag-mark 25 mm. wide by
2 mm. deep, tlanked on either side by intermittent shallow
scratches. This might be interpreted as a tail-trace but is more
probably the drag-mark of a Triassic conifer or horsetail rush.
Very similar marks in the Chirotherium sandstone of Thiiringeu
have been described as drag-marks of Equisetites by Riihle von
Lilienstern (1939, p. 370, pi. 12). Narrow shrinkage cracks,
rarely more than 5 mm. wide, characterize Level A.

452 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Level B, which carries the larger faunule, somewhat resembles
A in lithology. The sandstone overlying the recording surface,
however, quarries out in layered flagstones 1.5 to 3.5 em. thick;
the flat upper surfaces typically bear shallow ripples and run-off
marks. Shrinkage cracks of variable width are common. Rain-
drops have pitted both ground surface and footprints, obscur-
ing the plantar surfaces; but the sides of several deeper tracks
show striated claw-marks and fringe-scale furrows. Certain
parallel series of sharp, shallow scratches may be of arthropod
origin, but no trackway sequences are preserved. Annelids have
perforated the layers after deposition ; their sand-filled burrows
loop down into the shale or course along the sandstone-shale
interface, looking superficially like dasycladacean algae. To the
Milford flora recorded by Bock (1952, 1952 A) may be added
the genus Neocalaniites (MCZ 211). Several slabs bear branch-
ing or reticular patches, presumably algal.

The third Milford footprint horizon, discussed in my previous
paper (1954), lies in a reddish-brown siltstoue which crops out
in the Smith Clark quarry some 15 meters above Level B. Rep-
tile footprints of pseudosuchian appearance occur at another
horizon in that quarry but are unfortunately too vague and
fragmentary for adequate characterization (see Bock, 1952, pi
49, fig. 8).

The stratigraphic correlation of these beds with those of
southeastern Pennsylvania, the Connecticut Valley, and central
Europe will be discussed after an analysis of the fauna.

SYSTEMATIC DESCRIPTIONS

Nineteenth-century publications on the Milford footprints are
so vague, confused, and contradictory that little faith can be
placed in them. The foundations of Triassic ichnology had been
so monumentally laid by Edward Hitchcock that footprints from
any part of the Newark series were customarily identified by
reference to Hitchcock's figures of species from the Connecticut
River Valley. This pi'actice led inevitably to confusion, for the
classic Connecticut Valley faunas actually have little in com-
mon with those of Pennsylvania and New Jersey. As the nomen-
clature of Triassic footprints evolved, faunal revisers changed
the names of Milford species without checking the determina-

TRIASSIC REPTILE FOOTPRINTS 453

tions. For brevity, the terminologies of earlier authors and of
this paper are summarized in Table 1.

Before Bock's restudy, only C. H. Hitchcock (1889) had
erected new species for Milford footprints. His descriptions,
unaccompanied by figures, are so vague that of the species
mentioned only one, Chirotherium ["Otozoum"] parvum, can
be identified positively. His new species "Chiniaerichnus in-
gens," characterized only as "two relief tracks of considerable
size and representing only one-half of the foot," is a nomen
nudum. Any nomenclatural questions arising from C. H. Hitch-
cock 's work have, fortunately, been settled by Bock 's designation
of types.

Order SAURISCHIA

Suborder THEROPODA

Infraorder COELUROSAUBIA

Form-family GRALLATORIDAE Lull, 1904

(==ANCHISAURIPODIDAE Lull, 1904)
Genus CtRALLATOR E. Hitchcock, 1858

Type species G. cursorius E. Hitchcock (1858, p. 72) designated
by Lull (1904, p. 494).

Grallator sulcatus Baird, n. sp.
Figure 1 ; Plate 1, figure 1.

Type. MCZ 215, a left pes imprint.

Hypodigm. MCZ 215-228 inclusive, slabs bearing 37 pes im-
prints; AMNH 1982 and 1983, 6 imprints; LC S487, parts of
3 overlapping imprints. All collected by John Eyerman, 1885-
1887.

Horizon. Upper Triassic (Middle Keuper), Newark series,
upper Brunswick formation. Level A.

Locality {-fide Eyerman, 1889, p. 32). Quarry of the Messrs.
Clark about one-half mile east of the Smith Clark quarry, near
Milford, Hunterdon County, New Jersey.

454

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

TABLE 1 : ATTEMPTED SYNONYMY OF MILFORD FOOTPRINTS
Dashes indicate derivation from a previous author.

EYERMAN, 1886

C. H. HITCHCOCK, 1889

EYERMAN, 1889

LEVEL A :

aff. Anomoepus major

Brontozoum isodaetylum
Grallator cuneatus

[ ?] Grallator parallelus

[?] Grallator cuneatus

[ ?] Grallator cuneatus

[?] Grallator gracilis

LEVEL B :

aff. Anomoepus major

Otozoum parvum, n. sp.

[ ?] Grallator tenuis

Anomoepus minor

[?] Chimaeriehnus ingens, n. sp.

[?] Polemarchus gigas

Argozoum dispari-digitatum

Diagnosis. A moderately large Grallator, differing from other
species in the anterior position and relative shortness of digit

III and in having the bases of II and III closely united, while

IV is separated from III by a pronounced sulcus which extends
back to the metatarso-phalangeal pad of III. Ovoid metatarso-
phalangeal pads of II and IV regularly and equally impressing,
that of II more anterior. Second phalangeal pad of II opposite
first pad of III. Trackway characters unknown.

Measurements. Defects of impression and preservation make
impossible a valid and consistent suite of measurements. Dimen-
sions of the best-preserved specimens, however, suffice for de-
scriptive if not for statistical treatment. Measurements of the
type, adjusted for slippage and a transverse crack-filling, are
as follows :

TRIASSIC REPTILE FOOTPRINTS

455

LULL, 1915

BOCK, 1952

BAIRD, 1957

Anchisauripus parallelus
Grallator cuneatus — —

Grallator

' Grallator sulcatus, n.sp.

Grallator gracilis

♦Chirotherium parvum

Sauropus barrattii

Sauropus ingens
Polemarchus polemarchius
Argoides macrodactj'lus

Gigandipus ? (Anchisauri-
pus) milfordensis, n.sp.
Grallator gracilis

Chirotherium parvum
Chirotherium copei, n.sp.

Otozoum (?) lineatus, n.sp.
Sauropus barrattii

. Anchisauripus milfordensis

Anchisauripus parallelus
Genus incertum

Chirotherium parvum
Chirotherium eyermani, n.sp.

> Apatopus lineatus, n.gen.

(Unidentifiable)
Ehynchosauroides hyperbates,
n.sp.

Pes, digit no. H HI IV

Length, mm. 54 72 59

Length including metatarso-phalangeal pad 78 98 84

Length of pes, 100 mm.; including metatarso-phalangeal pads,

123 mm. ; width, 60 mm.

Measurements of digit divarication will vary greatly in a
single footprint depending on one 's reference points. In the type
specimen the angles between axes drawn through the centers
of claw base and metatarso-phalangeal pad are : II-7°-III-19°-IV.
Grallator sulcatus footprints, however, show quite a range of
divarication values. In my experience linear and angular meas-
urements such as these have a very limited value for the diag-
nosis and differentiation of theropod footprints.

456

BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY

Morphology. The abundance of deeply-impressed footprints
of this species affords an unusually clear picture of its foot
structure, including the relationship of the distal ends of the
metatarsals which is rarely recorded in Grallator trackways.
As in other theropod dinosaurs and in most cursorial birds the

Fig. 1. Grallator sulcatus n. sp., x %. Composite outline based on the
four best-preserved specimens. Section A- A' across the inverted natural cast
shows the close association of digits II and III and the sulcus separating
digit IV. Proximal phalanges of the restored skeleton appear foreshortened.

gait was digitigrade with most of the body weight falling on the
joints between the first and second phalanges of digits II, III,
and IV. Nevertheless, the pads which underlay the joints be-
tween metatarsals and phalanges of digits II and IV impressed

TRIASSIC REPTILE FOOTPRINTS 457

regularly, though less deeply than the digital pads, even in
shallow imprints. Apparently this Brunswick species had not
achieved the advanced degree of digitigrady shown by later in-
habitants of the Connecticut Valley. Unfortunately the lack of
trackway measurements precludes any useful comparison be-
tween gait patterns of Brunswick and Connecticut Valley
Grallatoridae.

The claws of Grallator sulcatus are acuminate in some indi-
viduals but blunted by wear in others. Regardless of depth of
impression the claws were always extended, rather than flexed
as in the other Milf ord theropods ; in particularly deep tracks the
exaggerated depth of claw-marks is due not to flexure but to
pivotal down-turning as the posterior part of the foot was raised.

The slightly domed phalangeal pads coalesce to form a sole
callus w^hich appears to have been slightly wider than the toe
proper, as in many cursorial birds. The three metatarso-phalan-
geal pads, though they impressed less deeply, form an integral
part of the sole and are separated from the phalangeal pads only
by shallow sulci. The close union of digits II and III and the
broad separation of IV differentiate Grallator sulcatus strikingly
from its congeners, in which the digits are either laterally
equidistant or III and IV are more closely appressed than II
and III.

Variation. Footprints from Level A reveal a sobering degree
of variability both in apparent form and in manner of impres-
sion. An increased divergence of the lateral toes is less common
than an exaggerated out-turning of the claws alone. Whether
this out-turning reflects some lateral flexibility at the ungual
joint, or merely individual variation in the plane of a hori-
zontally immobile joint, cannot be determined without trackways
showing successive imprints of the same foot. Distorted deep
impressions reveal that the lateral toes were spread apart as the
foot sank into the mud and were contracted again in birdlike
fashion as the foot was withdrawn.

Many of the footprints are so deformed by accidents of im-
pression that they give a decidedly erroneous picture of the foot
structure. Such anomalies serve to emphasize the dangers
involved in any attempt to characterize and interpret footprint
species without adequate quantities of well-preserved material.

458 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

A footprint is not an organism but the by-product of dynamic
contact between an organism and its environment. The all-too-
common typological analysis of isolated examples cannot be
expected to yield zoologically significant information.

Osteology. Heilmann (1927, p. 179 fP.) and Peabody (1948,
p. 399 ff.) have independently concluded that in the Grallator-
idae the phalangeal pads must have underlain the joints. This
is the situation in many cursorial birds, in which only the ungual
joint may coincide with a transverse crease. The osteological
restoration of Grallator sulcatus in Figure 1 conforms to this
principle but differs enough from the examples cited by Heil-
mann and Peabody to require explanation. In my opinion the
pes of Procompsognathus which Heilmann (1927, fig. 130 K)
has superimposed on the footprint of Grallator tenuis would fit
a great deal better if the foot were advanced until the ungual
joints of digits II and III corresponded with the distal pads in-
stead of their creases. All the joints would then line up neatly
opposite pads, with both ends of the fourth phalanx of digit IV
included in the distal pad. Except for the over-long claws of
Procoynpsognathus, foot and footprint correspond well in struc-
tural pattern.

The otherwise convincing skeletal restoration of Grallator cur-
sorius in Peabody 's figure 37 B shows a crease coinciding with
the joint between phalanges 3 and 4 of digit IV. Here again I
would prefer to enclose both ends of phalanx 4 in the distal
pad: and this is readily done if we use Hitchcock's original
figure of the footprint (1858, pi. 13, fig. 3), in which the distal
pad appears longer and more square-shouldered like that of G.
tenuis. Other Triassic theropod footprints such as Anchisauri-
pus and Eubrontes definitely have a long distal pad on digit IV,
and an occasional median constriction of this pad suggests that
two joints are actually represented. When the bones of any of
these footprint types are restored on the plan advocated here,
the phalangeal pattern is precisely that of Triassic and later
theropods : the phalanges of digit IV are short, stout-shafted, and
successively decreasing slightly in size.

No trace of a hallux is visible in any Grallator sulcatus foot-
print even where the metatarso-phalangeal pad of digit II is
impressed as deeply as 14 mm. Obviously the hallux, while pre-
sumably present, was vestigial and completely non-functional.

TBIASSIC REPTILE FOOTPRINTS 459

The lengths of the proximal phalanges, usually indeterminate
in more digitigrade coelurosaur footprints, can be estimated with
fair accuracy from the positions of the regularly impressed
metatarso-phalangeal pads. Pads II and IV are impressed to
equal depths, differing only as one side of the footprint is deeper
than the other. This situation contrasts sharply with that in
other species of Grallaior and in Anchisanripns and some species
of Euhrontes, where the metatarso-phalangeal pad of digit II
impresses less deeply or not at all. From this anomaly we may
obtain some clue to the probable structure of the metatarsus.

If a Newarkian coelurosaur such as Podokesaurus is restored
in a walking pose based on Grallator trackways (cf. Von Engeln
and Caster, 1952, fig. 277), the metatarsus slants forward and
downward at an angle of some 115° to the ground plane as
the foot is implanted. As the metatarso-phalangeal pads were
evidently developed to cushion the sole against the thrust trans-
mitted down the metatarsus, and as they served this function
only during implantation and the beginning of the propulsion
phase, it follows that the position of their imprints is (broadly
speaking) that of projections of the cross-sectional areas of their
respective metatarsals when in the implantation position. If
this be the case, as comparison with cursorial birds suggests, then
the type of transverse arching of the metatarsal bundle at its
distal end determines whether pad II or pad IV, or neither,
makes the more posterior impression. Similarly, the relative
length of the lateral metatarsals plus the thickness of their
underlying pads determines the relative depth of the pad im-
pressions. Of course the relative thickness of the pads is un-
knowable, but where the imprint of pad II is shallow or absent
(as in most Newarkian theropod footprints) metatarsal II was
probably shorter than IV. Conversely, the dinosaur represented
by Grallator sulcatus appears to have had lateral metatarsals of
approximately equal length.

Among Upper Triassic coelurosaurs metatarsal II is decidedly
shorter than IV in Procompsognathus , but the lateral metatarsals
are equal or subequal in Ammosaurus, Halticosaurus and Coelo-
physis. For comparison with Grallator sulcatus let us restore
from their pedal skeletons the footprints that would have been
made by two of these coelurosaurs.

460

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Halticosaurus liliensterni, a massive species from the KnoUen-
mergel of Thiiringen, is described by Huene (1934) as having
lengths of 20.5, 22, and 20 cm. for metatarsals II, III, and IV;
digit I is unknown. To show the relative positions of the digits
in walking pose we must make a three-dimensional reconstruc-
tion of the pes; when this is projected onto a horizontal plane

•- i

Fig. 2. Eeconstructed right pedes of Keuper coelurosaurs, with restored
footprints. A. Ealtioosaurus, x %,. B. Prccompsognathus, x 1. Data from
Huene.

(analogous to a footprint-recording surface) the metatarsals
and proximal phalanges are properly foreshortened (Figure
2 A). Structures in the metatarso-phalangeal region indicate a
foot with a moderately compact base and strongly divergent
digits; the bulk of the animal and the proportions of the meta-
tarsus suggest that the posterior pads impressed regularly and
to nearly equal depths. The positions of the plantar pads are
deduced from the skeleton and a plausible reconstruction of their

TRIASSIC REPTILE FOOTPRINTS 461

original form can be derived from Connecticut Valley footprints
of equivalent size.

The footprint thus restored is certainly neither a Grallator
nor a Gigandipus, and apparently not an Anchisauripm sensu
stricto. In Eiihrontes giganteus and Anchisauripus minusculus
we find the closest similarities in size, relative lengths of digits
and phalanges, and regular and subequal imprints of the meta-
tarso-phalangeal pads. Without venturing any positive correla-
tion we can say that these footprints may well have been made
by coelurosaurs of the Halticosmirus type.

Procompsognathus friassicus from the Stubensandstein of
Wiirttemberg is, in contrast to the hulking Halticosaurus, a
graceful and diminutive species. When reconstructed (from
Huene, 1921) and projected as before (Figure 2 B), its foot ap-
pears strikingly slender and laterally compressed. Neither the
hallux nor metatarso-phalangeal pad II normally made contact
with the ground. The restored footprint is obviously that of a
Grallator closely comparable to a small G. cursorius but having
a relatively shorter central digit. If the divarication were
larger and the lateral digits somewhat shorter, correlation with
a small G. tenuis would be indicated.

A comparison of Figures 1 and 2 shows clearly that, even dis-
regarding size, Grallator sulcatus has much stronger resem-
blances to Procompsognathus than to Halticosaurus. So far as
osteology can be inferred from the footprint, only size, minor
differences in digital proportion, and the shortness of metatarsal
II distinguish Procompsognathus from the Milford trackmaker.

Unpublished data on skeletons of Coelophysis from the Upper
Triassic of New Mexico, made available for this study through
the courtesy of E. H. Colbert, show that the foot of that genus
is essentially similar to that of Procompsognathus except that
the lateral metatarsals are equal in length. Coelophysis thus
closely approximates in structure the reconstructed skeleton of
Grallator sulcatus. While no positive correlation can be made,
the Level A footprints attest the presence of a Coelophy sis-like
coelurosaur in the Newark basin of deposition.

462 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

Form-Family GRALLATORIDAE Lull, 1904

Genus AnCHISAIIRIPUS Lull, 1904

Type species A. sillimani (E. Hitchcock), 1843.

Anchisauripus milfordensis (Bock)
Figure 3 A

Gigandipus ? (AncMsaiiripiis) milfordensis Bock, 1952, p. 403 ; pi. 43, fig.
3 (in legend, for "No. 15210" read S488).

Type. LC S488, a left pes imprint slightly distorted by the
paratype pes of Chirotherium parvum.

Hypodigm. The type and two other imprints on the same
slab; MCZ 135 and 229; AMNH 1981 and 1984. These seven
imprints collected by John Eyerman, 1885-1887.

Horizon. Upper Triassic (Middle Keuper), Newark series, up-
per Brunswick formation about 5100 meters above the Triassic
base. Level B.

Locality. Smith Clark quarry near Milford, Hunterdon
County, New Jersey, 1.25 km. north of Delaware River bridge.

Tentatively assigned. Certain tracks from the Gettysburg shale
(a Brunswick equivalent) near Goldsboro, York County, Pennsyl-
vania (Wanner, 1889, pis. 6, 7, 10). This material was acces-
sioned by the U. S. National Museum in 1888 but is now lost,
so that precise comparisons are impossible.

Diagnosis. A small, short-footed, broad-based species. The
base of claw II lies opposite the crease between the first and
second phalangeal pads of digit III, and the tip of claw IV
opposite the distal part of the second pad. Metatarso-phalangeal
pads II and IV circular and strongly domed, IV nearly twice
the size of II. Claws habitually flexed, hastate in outline with
basal recesses.

Morphology. Collation of the new hypodigm permits some am-
plification and emendation of Bock's description. Removal of
some of the obscuring underclay reveals that the structure which
Bock interpreted as a semi-rotated hallux is actually a filled
shrinkage-crack which runs from the margin of the slab through
metatarso-phalangeal pad II of the type, narrows abruptly, and
passes under the label and into the manus of Chirotherium
parvum. Nowhere is there evidence of a hallux imprint, even

TRIASSIC REPTILE FOOTPRINTS

463

in MCZ 135 where the first phalanoeal pad of digit II impressed
to a depth of 11 mm. This fact and the osteological differences
listed in Table 2 preclude assignment of the species to Gigandi-
pus.

Individual variation is apparent in even this small sample.
MCZ 135 in particular differs from the others in its relatively
shorter digit IV, its large size (103 mm. long vs. 93 mm. in

Cm.

I—

5

H

Fig. 3. A. Anchisauripus milfordensis (Bock), x 3/4. Composite outline
based on all seven Milford specimens, drawn the size of S488 or MCZ 229.
Section A-A' shows the domed lateral metatarso-phalangeal pads. B. Anchi-
sauripus parallelus (Hitchcock), AjMNH 1789, x %.

the smallest individual, the type), and its angles of digit divari-
cation (II-13°-III-10°-IV vs. II-15°-III-9°-IV in the type).

The outstanding features of the species — the broad-based
foot, the broad, rounded phalangeal pads, and the flexed, hastate
claws — have been adequately discussed by Bock. Metatarso-
phalangeal pad III is ill-defined, but the dome-like lateral pads
are impressed to approximately equal depths with reference to

464 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the rest of the foot. The lack of creases between the phalangeal
pads suggests the thick, relatively inflexible plantar padding of
a foot which is so specialized for cursorial habits that it retains
very little power of grasping. Fringe scales on the median side
of digit III are represented by parallel striae, somewhat irregu-
larly spaced, which average 10 to the centimeter. They tell little
about the squamation but at least prove that the full width of
the digit is recorded.

Osteology. A close comparison of Figures 1 and 3 A reveals
significant differences in the restored skeletal structure. The
individual phalanges of Anchisaurijnis milfordensis are rela-
tively shorter than those of Grallator sulcatus, and the unguals
may have been, like the claws, shorter and blunter. More im-
portant is the dift'ereuee in the relative positions of the digits.
While in G. sulcatus the joint of ungual IV lies opposite the
joint between phalanges 1 and 2 of diigt III, in A. milfordensis
it lies opposite the waist of phalanx III-2, and the joint between
phalanges I and 2 of digit III is aligned with the waist of
phalanx IV-3. Clearly the central digit of A. milfordensis is less
advanced with respect to the lateral digits. Additional osteo-
logical criteria for comparing this species to other dinosaur foot-
prints will be discussed in a subsequent section.

Anchisauripus parallelus (E. Hitchcock)
Figure 3 B

Material. AMNH 1789, a sharp impression of a right pes
associated with a footprint of Ajjatopus Uneatus. Collected by
John Eyerman, evidently in the Smith Clark quarry.

Discussion. In its form and proportions, especially in the
slenderness and lack of divergence of digit IV and the relative
positions of the articular pads, this specimen is indistinguish-
able from footprints of A. parallelus found in the Portland
formation of Massachusetts and Connecticut. Its reconstructed
skeleton corresponds exactly to that of the type specimen as
figured by Lull (1953, fig. 42). The Milford footprint, however,
is only 6/11 the size of the Massachusetts type. This striking
difference in size is associated with no significant difference in
proportions : one specimen is a miniature of the other. Despite

TRIASSIC REPTILE FOOTPRINTS 465

our ignorance of trackway characters and range of variation in
the Milford form, its form-specific identity is assured.

This is the only footprint species common to the Milford and
Connecticut Valley faunules. Whether it represents the same
reptilian genus in both areas is of course unknown.

Although no hallux impression is present on the slab, hallux
function within populations (or even individual trackways) of
Anchisau7-ipus is too variable to be of significance in an isolated
footprint. Deep creases separate the pads of digits II and III
and mark the bases of the strongly flexed lateral claws. Evi-
dently the foot of this dinosaur was rather flexible — perhaps
prehensile — at least more so than those of A. milfordensis and
Grallator sulcatus. The fourth digit of A. parallelus is longer
but clearly less robust than the second, and (at least in the
Milford form) appears to be closely joined to the strong third
digit. In this respect A. parallelus differs from A. milfordensis,
in which the lateral digits are about equal in robustness and in
independence of the third, and differs even more from G.
sulcatus, in which the second and third digits are joined and
the fourth exceptionally independent. The adaptive significance
and evolution of these structural patterns can, unfortunately,
only be surmised until stratigraphic sequences have been estab-
lished and skeletal material is available.

GENUS INCERTUM
Figure 4 D

A fourth and most peculiar form of dinosaur is represented
by a single small, deep imprint of a left pes, MCZ 214. This
footprint might be dismissed as an anomaly of impression if a
similar specimen had not been found by Wanner (1889) in the
Gettysburg shale of York County, Pennsylvania. Wanner 's
drawing is reproduced as Figure 4 E ; the loss of his original
at the U. S. National Museum precludes further comparison.

This animal had a broad, compact foot with the central digit
projecting only a toe-width beyond the lateral ones. The im-
print is deepest on the lateral side (where it is obscured by
worm burrows) and in the free lengths of the second and third
digits; the rest of the broad "sole" is two to three millimeters
shallower. Little rotational or lateral slippage seems to have

4(36 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY

Fig. 4. A-C, three dinosaur footprints drawn to common size and orienta-
tion, with phalanges restored. A. So-called Grallator gracilis C. H. Hitch-
cock, Dartmouth 5023. B. Type of Anchisauripus hitchcocki Lull, Amherst
56/1. C. Plesiotype of Grallator tenuis E. Hitc:hcock, Amherst 17/4. B and
C from Lull. One centimeter scale.

D. Tentative reconstruction of genus incertum from Milford, MCZ 214,
.X 1. E. Similar footprint from Goldsboro, Pennsylvania, x 1 (from Wanner).
Apparent sharpness of claw tips may be caused by shrinkage cracks.

TRIASSIC REPTILE FOOTPRINTS 467

occurred: the peculiar proportions of the foot are real. But the
material available is insufficient for proper analysis and certainly
inadequate for taxonomic treatment.

TAXONOMY OF THEROPOD FOOTPRINTS

Up to this point we have assigned dinosaur footprints from
the Delaware Valley to genera best represented in the Con-
necticut Valley, employing a necessarily typological classifica-
tion based on similarities and differences in footprint form. This
is the traditional approach pioneered by Edward Hitchcock in
his "Ornithichnology" of 1836. In Hitchcock's subsequent
papers an increasing number of form-taxa were variously
grouped and regrouped, with unfortunate wholesale renaming of
many units. In this century Lull has achieved a logical taxo-
nomic and nomenclatural reorganization of the Connecticut Val-
ley footprints, a classification which is crystallized in his recent
revision (1953).

The three determinable species of theropod footprints from
Milford are here assigned to two New England genera which
Lull has made the types of two families. Lull's family diagnoses
are not, however, mutually exclusive :

Grallatoridae : typically small, footprint tridactyl, limbs very
long ; with or without tail trace.

Anchisauripodidae : bipedal, tetradactyl; hallux when im-
pressed rotated to the rear; well marked phalangeal pads; an-
terior claws acuminate but not strongly raptorial ; no caudal
impression.

In current practice GraUator is distinguished from Anchi-
sauripus "by greater relative length of stride, smallness of track,
and the absence of a hallux impression" (Lull, 1953, p. 153).
Within each genus the smaller species are distinguished by size,
relative length of digits and of stride, and divarication of the
lateral digits. But intermediate forms are common, and isolated
or incomplete footprints may be difficult to assign to genus and
species even when they are clearly impressed.

Let us examine a specific example of close resemblance. A
trackway identified by C. H. Hitchcock as his GraUator gracilis
(Dartmouth 5023) and the type specimen of Anchisauripus
hitchcocki Lull (Amherst 56/1) are morphologically so similar

468 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

that the differences between them fall within the range of in-
dividual variation. Lull's plesiotype of Grallator tenuis E.
Hitchcock (Amherst 17/4) differs from them chiefly in having
digit III slightly farther in advance of the lateral toes (see
Figure 4 A-C). All are from the Portland formation of the
Turners Falls area, Massachusetts. Their differences in size,
gait, and incidence of hallux impression are tabulated below.
(Data on the Amherst specimens from Lull.)

"G. gracilis"
Dartmouth 5023

A. hitchcocki
Amherst .56/1

G. tenuis
Amherst 17/4

Pes length, mm.

51

119

73

Pace length

299

360 to 500

195

Pes : pace length

1:5.9

av. 1:3.6

1:2.7
(1:3.3 in type)

Hallux imprint

absent ( ? )

present

absent

What do these differences mean? Size difference may be
genetic or ontogenetic. Biometric analyses of true populations
of Connecticut Valley footprints have not yet been made, but
useful series of measurements from footprints of varied ages
and sources have been compiled by Lull (1953). In general,
dimensions in each species tend to cluster about a mean, with
"occasional gradational footprints which bridge the dimensional
gaps between the species. ' ' This has been interpreted as evidence
of several species of reptiles; but as Lull observes, differences
caused by sexual dimetry or by selective representation of age
groups (perhaps caused by seasonal migration) are difficult if
not impossible to evaluate.

The presence or absence of a hallux imprint appears to be a
valid distinction in the huge, broad-soled dinosaur tracks of
the Connecticut Valley : in Euhrontes the hallux is almost never
in evidence, while in Gigandipus it usually impresses its entire
length. In the Grallator-Anchisauripus group, however, the dis-
tinction is less clear. Rarely does more than the tip of the hallux
claw impress, and even this may be totally absent, as in a four-
imprint trackway of A. sillimani from Turners Falls (MCZ 252.
ex Dartmouth 14, cited in Grier, 1927). In current taxonomic
practice the absence of a hallux impression does not debar a
specimen from Anchisauripus but the presence of one debars it
fi'ora Grallator.

TRIASSIC REPTILE FOOTPRINTS 469

This distinction is unconvincing, especially when we consider
that the hallnx imprint is typically absent in the smaller and
present in the larger members of a morphologically homogeneous
series. The foot of a large individual bears relatively more
weight than that of a small one, as body bulk is proportional
to the cube of the linear dimensions: in walking this greater
weight on the metatarsus might well cause the pendant hallux
claw to impress more fretiuently. Similarly, a juvenile dinosaur
might be expected to take a relatively longer stride than its
more ponderous parent. For these reasons it seem unrealistic
to maintain generic, much less familial, distinctions between
dinosaur footprint species solely on the basis of such largely
extramorphologic characters as size, incidence of hallux imprint,
and length of stride.

/ subynit that the characters most diagnostic for the classifica-
tion of footprints as such, as well as most useful for comparison
with skeletal remains, are those which reflect the bony structure
of the foot. In most adequately-known varieties of dinosaur
footprints the presence of articular swellings and pads permits
a reasonably accurate analysis of the skeletal pattern.

To clarify the relationships of the Milford species, therefore,
let us compare their restored pedal skeletons with those of other
Upper Triassic theropod tracks. For this purpose we must
add the phalanges to published figures of the various species,
extrapolating to find the length of phalanx III-l where the posi-
tion of its proximal end is not indicated by a pad. Because of
possible errors in the interpretation of joint position in published
figures, this analysis should be considered provisional.

When the skeletons thus reconstructed are compared, the main
differences between them are seen to lie in (1) the relative
positions of the metatarso-phalangeal joints;^ (2, 3) the projec-
tion of the central digit beyond the lateral ones; and (4) whether
digit II or IV projects farther forward. These factors are of
course to some extent interrelated. They are more specifically
keyed out below, and their distribution among a number of foot-
print species is shown in Table 2.

1 This character, the most useful for classification and for skeletal correlations,
Is also the most difficult to determine in published drawings and faintly-impressed
tracks. Close analysis of original matorial will doubtless necessitate refinements
In the subdivision attempted below.

470 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

This method of analysis parallels in part the highly instructive
comparisons by means of Cartesian diagrams which Lull has ap-
pended to his 1953 monograph. (The new insights provided by
these diagrams have not, curiously enough, caused Lull to modify
his classification of 1915.) My approach, however, concentrates
on differences in presumed skeletal structure. By comparing
the species of several genera on a single objective basis, rather
than matching the previously assigned species of each genus
against the type species, it brings out striking inter-generic
similarities which might otherwise be overlooked.

DIAGNOSTIC CHARACTERS

1. A. Metatarso-plialangeal joint III about opposite waist of phalanx II-l

and joint 1-2 of IV.

B. Metatarso-plialangeal joint III about opposite proximal end of II-I
and waist of IV-1.

C. Metatarso-phalangeal joint III about opposite that of II and joint
1-2 of IV.

2. A. Joint of ungual II about opposite joint 1-2 of III.

B. Joint of ungual II about opposite waist of III-2.

C. Joint of ungual II about opposite joint 2-3 of III.

3. A. Joint 1-2 of III about opposite joint of ungual IV.
B. Joint 1-2 of III about opposite joint 3-4 of IV.

(\ Joint 1-2 of III about opposite joint 2-3 of IV.

4. A. Digit II projects farther anterior than IV.

B. Digits II and IV project about equally far.

C. Digit IV projects farther anterior than II.

Let US now attempt a generic regrouping of these species :
I. Grallator, type species G. ciirsorius. G. sulcatus clearly
belongs to this group despite its atypical forward-set central
digit and subequally projecting lateral digits. G. tenuis appears
to be intermediate between groups I and II, resembling the latter
especially in the relative positions of the metatarso-phalangeal
joints. A surprise here is that Otouphepus minor, though super-
ficially similar to 0. niagnificus, is otherwise quite unlike it and
shows itself to be a Grallator- in size and morphology. This
species, based on a single isolated footprint, is of questionable
validity.' G. forynosus, included doubtfully by Lull because of
its long stride and lack of a hallux imprint, belongs rather to
Group III.

- TliP nature of Otoiiiihc/iiix is discussed in Appendix 1.

TRIASSIC REI'TILE FOOTPRINTS

471

TABLE 2: COMPARISON OF SOME UPPER
TRIASSIC THEROPOD FOOTPRINTS

1.

2_

3.

4.

Grallator cursorius

A

A-B

B

A

O G. cuneatus

A

*A

B

A

~ G. gracilis-''

A

A

B

f

2 Otouphepus minor

A

A

B

B

- G. sulcatus (Figure 1)

A

A

A

B

G. tenuis (Figure 4 C)

B

A-B

B

B

O Anehisauripus sillimani

B

B

B

A

O "A. exsertus" plesiotype, Amherst 54/1

B

B

B

A

i2 A. hitchcoeki (Figure 4 B)

B

A-B

B

B

- Dartmouth 5023 (Figure 4 A)

B

A-B

B

B

Anehisauripus tuberosus

B

A-B

C

C

A. exsertus

B

B

C

C

- A. parallelus (Figure 3 B)

B

B

C

C

S A. australis (Argentina)

B

B

C

C

2 Jeholosauripus ssatoi (Manchuria)

B

B

C

B-C

S, Grallator formosus

B

A-B

C

C

- Otouphepus magnificus

?B

B

B-C

B

A. minusculus

B

A-B

C

B

A. milfordensis (Figure 3 A)

B

A

B-C

B

Eubrontes giganteus

B

B

C

B

Gigandipus caudatus

C

C

B

A

II. Anchisauripus (sensu stricto). A. sillimani, the type
species, shows closest affinities with A. hitchcoeki, G. tenuis, and
the Dartmouth trackway assigned to G. gracilis. Amherst speci-
men 54/1, figured by Lull (1904, fig. 7; 1915, fig. 41; 1953, pi.
10, fig. 11 — but not fig. 41 which is mis-captioned Amherst 54/1
but actually shows 16/6, the type) as Anchisauripus exsertus
proves instead to be a large A. sillimani. (The same is true of
Amherst 34/14 in Lull's pi. 10, fig. 12.) Much original material
of these forms must be compared before taxonomic decisions can
be reached.

ii Lull's figure of the O. gracilis plesiotype appears rather different from C. H.
Hitchcock's original tigure of the type (in E. Hitchcock. 1805, pi. 9, fig. 7).
Perhaps the latter figure is inaccurate or the plesiotype is misassigned.

472 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

III. Anchisaurepus (sensu lato). Quite a distinct aspect is
shown by the larger species of Anchisauripus (earlier assigned to
Brontozoum) , in which digit IV is set well forward. Within
this group A. tuberosus, A. exsertus and A. parallelus are so
similar that they might be considered a single variable species.
A. australis Lull (1942) from the Pagazano beds of Argentina
is a typical representative of the group. Jeholosauripus ssatoi
from the ''Eo-Mesozoic" of Manchuria (Shikama, 1942) clearly
belongs here, as do Grallator formosus and Otouphepus mag-
nificus. Suppression of the latter genus and species (Baird,
1956; see Appendix I) will remove them from consideration.
A. minusculus is, as Lull notes, a sharply defined species ; it
seems to be intermediate between the other members of this
group and Euhrontes giganteus. A. milfordensis is also some-
what transitional but is referred to Group III because of its
forward-set central digit and its small size.

Strict consistency would require that Group III be dis-
tinguished as a separate genus of the Grallatoridae. Unfortu-
nately there is no Hitchcockian name available for these forms,^
and rather than make major nomenclatural changes on the basis
of this preliminary survey, I will for the present follow Lull's
terminology.

The dinosaurian footprint associated with Chirotherium lulli
at Milford (Baird, 1954, fig. 2 C) and the specimen from York
County, Pennsylvania, which Hickok and Willard (1933, fig.
6 B) assigned to Anchisauripus sillimani are so different from
the forms in Table 2 (and from the Anomoepodidae as well)
in the position of their lateral digits that they cannot be classi-
fied in the key used here. They apparently belong to a distinct
family, which is represented in Europe by Coelurosaurichnus
(sensu stricto: see Baird, 1954, p. 182). This family is appar-
ently not represented in the Meriden and Portland formations
of the Connecticut Valley.

4 The appropriate ami previously used name Brontozoum is excluded by nomen-
clatural technicalities which are discussed iu Appendi.x IT.

TRIASSIC REPTILE FOOTPRINTS 473

Order THECODONTIA

Suborder PSEUDOSUCHIA

Form-family CHIROTHERIIDAE Abel, 1935

Genus ChirOTHEKIUM Kaup, 1835

Brachychirotherian Group

Diagnosis. Specialized Upper Triassie chirotheriids lacking a
thumb-like fifth phalangeal segment distinct from the metatarso-
phalangeal pad.

Chirotherium parvum (C. H. Hitchcock), 1889
Figures 5, 6 A ; Plate 1, figure 2.

Otozoum parvum C. H. Hitchcock, 1889, pp. 122, 123, 127.

Oiozoum parvum. Lesley, J. P., 1889, pp. 571-573, 3 figs.

()tn~oum parvum. Lyman, B. S., 1894, p. 214; 1895, pi. 608, figs. 1-3.

'.Oinznum parvum. Lull, E. S., 1904, p. 515.

'.rhirnfherium parvum. Lull, E. S., 1915, p. 226, fig. 77.

Clieiroiherimn (?) parvum. Lull, E. S., 1917, p. 119.

rhirollnnium parvum. Peabody, P. E., 1948, p. 346.

nnroilurinm parvum. Bock, W., 1952, pp. 410-414; pi. 41 with plate title

42; pi. 42 with plate title 41; pi. 43, fig. 2 (for Paratype No. S488

read Ilolotype No. S490).
Chirotherium copei Bock, 1952, pp. 414-415; pi. 43, fig. 1 (for x 1 . . .

No. S491 read x 1/2 . . . ANS 15210).
Chirotherium [copeil. Eichards, H. G., 1953, fig. 176.
Chirotherium parvum, C. copei. Baird, D., 1954, pp. 174, 175.

Type. LC S490, designated by Bock, a large right manus-pes
set; the missing posterior half of metatarso-phalangeal pad V
is preserved on MCZ 212.

Hypodigm. The type, LC S488 and MCZ 211 probably rep-
resent two individuals of similar size. ANS 15210 (type of C.
copei), MCZ 209, MCZ 210 and AMNH 2257 appear to represent
one individual. Horizon, locality and collector as for Anchisauri-
pus milfordensis (collector of ANS 15210 is unknown).

Diagnosis. Phalanges of pes digit V reduced and included
in the metatarso-phalangeal pad ; pes digits IV and V clawless ;
narrow, curved claws on pes digits I to III borne high above the
thickly padded plantar surface and divergent laterally; meta-
tarso-phalangeal pads III and IV coalesced.

474 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

History. The type of this remarkable species has been validly
designated by Bock, yet curiously enough this may not be the pri-
mary specimen on which Hitchcock's species concept was based.
The original description, fragmented between two sections of a
notoriously imprecise paper, designated no type and was not
illustrated. Outline figures of the species, however, appeared the
same year in Lesley's hodgepodge "Dictionary of Fossils" wnth
this notation :

"Otozoum parvum, n. sp. C. H. Hitchcock, Fig. 1, hind

foot track; fig. 2, fore foot track (both natural size) ; fig. 3,

reduction of the two foot prints to show in what relation they

stand to each other on the slab of Trias sandstone in the

quarries at Milford, on the New Jersey side of the Delaware

river, about thirty miles above Trenton. Discovered and

traced iu outline by Prof. C. H. Hitchcock. (See MS. letter,

Dec. 1888.) Upper Trias. — Note. When first seen it was

thought to be a Cheirotherium track, but it has only four

toe marks on each foot." (Lesley, 1889, pp. 571-572.)

These figures, though reprinted by Lyman (loc. cit.), have

hitherto escaped the attention of bibliographers and researchers

alike.

A second outline drawing by Hitchcock, first published in
1915 by Lull, is not identical with the first. Both, however, show
the cast of a right manus-pes set, and they are so similar in
proportions and particularly in the relative positions of the
manus and pes (seldom duplicated exactly in a chirotheriid
trackway) that both probably represent the same specimen.

Only one of the existing specimens, the type of C. copei Bock,
closely resembles these drawings in the features cited. Hitch-
cock's statement that the pes "is 5 inches long besides 2y^
inches of heel" applies to this specimen but cannot refer to the
Lafayette College types. Now the early history of the C. copei
slab, which Bock obtained (mislabeled as to locality) from the
efl^ects of an amateur collector of fossils, is obscure ; yet the in-
ternal evidence indicates that this specimen or one remarkably
like it constituted the undesignated, conceptual type of Hitch-
cock's species. Thus Chirotherium copei may be an objective as
well as a subjective junior synonym of C. parvum.

Variation. The three individuals (or so) represented in the
population show a good deal of variation and (if the isolated

TRIASSIC REPTILE FOOTPRINTS

475

tracks are correctly assigned) some intra-individual variation
as well. These irregularities are, of course, in addition to those
caused by differences in impression and preservation. The type
and paratype may represent the left and right feet of the same
individual. Pes digit I of the type, however, is relatively longer :
its claw-mark extends nearly to the tip of digit II instead of to

Fig. 5. Chirotherium parvum (C. H. Hitchcock), right manus-pes sets,
X 1/3. A. Type of C. copei, ANS 15210. B. Type of C. parvum, 8490 +
MCZ 212, showing striated digit-tip impressions.

476 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

the base of the distal pad on II, as in S488, MCZ 210, and ANS
15210. MCZ 209 is intermediate in the length of digit I but is
otherwise nearly a mirror image of ANS 15210. In relative
width of the digit I-TV group and in robustness of manus digit
I the type and paratype resemble each other and differ from
the smaller, slenderer specimens which are referred to the "C.
copei" individual (cf. Figure 5).

In my opinion these differences are insufficient to distinguish
two species. Individual, age, and sexual variation are to be ex-
pected in the Milford reptile population. Differences observed
within the hypodigm are minor and do not appear to be corre-
lated. All the specimens exhibit a syndrome of specializations
which is unique within the form-genus Chirotherium and almost
certainly represents a single reptilian genus. Sympatric species
of this unusual genus are of course possible, but since similari-
ties greatly outweigh differences, the burden of proof would seem
to lie with the splitter.

Morphology. Most of the salient features of this species have
been described by Lull and Bock. Curiously enough the pes
claws, which are present in every specimen and constitute one
of the most distinctive features of the species, remain unde-
scribed and their existence has only been surmised (Bock, 1952,
p. 412). These claws were carried high above the plantar surface
so that only their tips ordinarily impressed. Spalling of the
natural casts obscures the relationships, but where measure-
ments can be made the base of the claw lies at least 6 mm. above
the sole of the digit tip.

Instead of forming a linear extension of the digit these claws
are turned strongly outward, so much so that the first and
second must have nearly touched the sides of the digits lateral
to them. The claws were carried in the normal vertical posi-
tion; their lateral divergence of some 35° seems to be the effect
of an oblique ungual articulation, for the digits themselves are
otherwise nearly straight.

Claw I is long and narrow and slightly curved laterally; its
distal half forms a convex ventral keel rather than a point. Claw
II, the heaviest, is carinate along its concave ventral profile but
ends in a conical point, sometimes blunted by wear. Claw III
is similarly curved and pointed. Of course the depth of the claw-
tip impressions has been exaggerated when the toes dug in at

TBIASSIC REPTILE FOOTPRINTS 477

the end of the propulsive phase. Digit IV appears to be claw-
less in all five specimens, even where the digit-tip impression
is 8 mm. deep — an anomalous condition, for a well-developed
I'law IV characterizes most other chirotheriids.

The vestigial fifth pes digit, so different from the well-de-
veloped "thumb" of most chirotheriids, is a striking feature of
Chirothcrium parvum. A single ovoid pad underlies the pha-
langes and the metatarso-phalangeal joint. As in several other
Keuper species (Baird, 1954, p. 174) this pad has migrated
somewhat medially from the primitive position, so that the distal
end of metatarsal V must have underlain the shaft of IV. Onl.v
slight marginal indentations and a suggestion of separate plan-
tar thickenings distinguish the phalangeal section of the pad,
which is scarcely longer than its width and narrows abruptly
to an ogival, clawless tip.

Almost as distinctive as this digital "heel" is the pattern
of plantar pads underlying the metatarsal cross-axis. The meta-
tarso-phalangeal pads form a straight line of low bosses across
the posterior edge of the sole ; a single flat, subeircular pad
underlies the bases of digits III and IV. This condition is quite
unusual among chirotheriids. A pad common to two digits occurs
in both large-manus and small-manus species but this pad is
always central, joining the bases of digits II and III, while the
pads of I and IV are distinct. Only in Otozoum do we find a
single pad for digits III and IV.

Unlike the pes, the manus is typically chirotherioid in form
and shows little specialization except that claws are apparently
absent and digit V is only slightly offset. (Digits IV and
V are definitely clawless but evidence on the others is incon-
clusive.) The fifth digit is not small and abortive, as Bock
terms it, but normal in length and robustness. In the exception-
ally deep paratype imprint, moreover, its pad is revealed as
merely the slenderer distal part of a large ovoid "heel" which
extends postero-medially to a point in line with the axis of digit
I. (See Bock, 1952, plate 41 with plate title 42.)

The imprint made by this "heel" pad is 13 mm, deep and
slightl}^ undercut laterally. Its steep posterior and postero-
medial margins are in strong contrast to the gently sloping mar-
gins of the pedal "heel" and suggest that the fifth metacarpal

478

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

cm.

10

I ''■■'■■'* I

cm.

10

1 1 1 1 1 1 1 1 1 1 1

TRIASSIC REPTILE FOOTPRINTS 479

was held nearly vertical as the manus was implanted. The other
metatarsals apparently sloped less steeply than the fifth but con-
siderably more steeply than the metatarsal bundle. This evidence
of a more digitigrade manus supports the correlation of Chiro-
therium parvum with a reptile of bipedal ancestry in which the
forelimbs were shorter than the hind, and in which such a func-
tional lengthening of the forelimb pendulum would have made
for more efficient quadrupedal locomotion.

The paratype manus imprint is divided by a large shrinkage
crack which, oddly enough, does not exaggerate its width but
rather compresses it laterally so that the third and fourth digit
impressions are coalesced. The type imprint of Anchisauripus
milfoi'densis is also compressed where it is crossed transversely
by the same crack. My explanation for this phenomenon is that
the shrinkage crack antedated the footprints (see Soergel, 1925,
figs. 45-48 for other examples). Before the sand which now
forms the natural casts was washed over the recording surface
the clay was flooded and consequently expanded, narrowing
the shrinkage cracks and thus compressing the footprints. Ich-
nologist, take warning: things are not necessarily what they
seem.

Chirotherium eyermani Baird, n. sp.
Figure 6 B ; Plate 2

Eyerman's chirotherium. Baird, 1954, pp. 174, 175-176.

Type. MCZ 134, an isolated left pes imprint collected by John
Eyerman in 1887 ; the only known specimen. Horizon and
locality as for Anchisauripus niilfordensis.

Diagnosis. A moderately large species with slender digit I,
short, robust pes digits II-IV bearing heavy claws at sole level,
and greatly shortened, hoof -like digit V enclosed in a single pad.
Digit IV shorter than II. Metatarsal cross-axis oblique; plantar
padding undifferentiated. Manus and trackway unknown.

Fig. 6. Brachychirotherian (A, B), small-manus (C), and large-manus
(D) chirotheriids with attempted skeletal restorations. A. C. parvum, com-
posite of entire hypodigm. B, C. eyermani n. sp., type (MCZ 134). C. C.
lomasi, Keuper of Storeton, Cheshire (Yale Peabody Museum 3762, a small
individual). D, C. barthii, Moenkopi of Cameron, Arizona (University
of California Museum of Paleontology 37315). A-C x 1/3, D x 1/5.

4v0 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

MorpJiology. The single footprint of this species, which is im-
pressed to a maximum depth of 40 mm. below ground level,
illustrates a fact often overlooked : a footprint is not a cast of
the underside of a foot but a composite record of the foot in
motion. The effects of structure, function, and preservation are
combined in the specimen and must be separated by analysis.
If we eliminate the functional distortions of the imprint (which
are revealed by slippage marks and variously inclined groups of
striations produced by the claws and scales) the foot itself
appears to have the form shown in Figure 6 B.

A tendency toward functional tridactyly is evidenced by a
strengthening of the three central digits at the expense of the
first and fifth. Indeed, the soft mud was resistant enough to
bend the feeble first digit upward at a 20° angle to the second.
The inward-turned imprint of claw I may be atypical, for such
a deflection occurs sporadically in chirotheriid traclcAvays. The
undulant outlines and almost undifferentiated pads of digits II-
IV are reminiscent of C. parvuni, but in strong contrast are the
straight, massive claws with their slightly convex soles forming
an unbroken continuation of the general plantar surface. Also
distinctive is the shorter, more divaricate digit I-IV group with
its more oblique metatarsal cross-axis and flat, subrectangular
sole. From the bases of digits I and IV a pair of ridges which
appear to represent the flcxores digitorum hreves extend prox-
imally, converging toward the tarsus. The slope of the longi-
tudinal arch, a feature rarely revealed in chirotheriid footprints,
indicates that the first four metatarsals formed an angle of 155°
with the proximal phalanges as the pes was implanted.

Digit V, revealed in unusual detail in this footprint, closely
resembles that of C. parvum except that its phalangeal segment
is less pointed and is quite undifferentiated from the metatarso-
phalangeal pad. This "heel" pad is relatively flat instead of
ovoid as in C. parvum; it is impressed about 9 mm. deeper than
the ventral surface of the longitudinal arch. A definite bridge
connecting the "heel" to the base of digit IV suggests a liga-
mentous connection between the distal ends of the metatarsals.
From the sole at the tip of digit V a small falciform web curves
upward and joins the side of digit IV, well above its sole.

TRIASSIC REPTILE FOOTPRINTS 481

The distal margin of digit V shows a striation of flutings
which average 2.5 mm. between crests. This fluting resembles
the striated 'impressions of claws III and IV and is quite distinct
from the scalloped profile produced by fringe scales 3 mm. wide
on the medial border of digit II. The greatly reduced fifth digit
thus appears to have been tipped with a broad nail or hoof.
Such a structure has been reported in no other species of Chiro-
therium.

Osteology of Level B Chirotheriids

The thick, little-difiPerentiated plantar padding in Chiro-
therium parvum and C. eyermani makes it impossible to locate
all the joints and thus to determine precisely the phalangeal
formula. Representative phalangeal patterns may, however, be
deduced in other species where articular swellings are well de-
veloped. Figure 6 D shows the arrangement in C. harthii, the
common large-manus species of the European and American
Lower (or Middle) Triassic. Here the formula indicated is
manus 2-3-4- ?4-?3, pes 2-3-4-5- ?4. An Upper Triassic member
of the same group, C. lulli, is basically similar but appears to
have only three phalanges in the thumb-like fifth digit (Baird,
1954, fig. 2 A). In the best-known small-manus species, C. lomasi
of the English Keuper (Figure 6 C), a formula of 2-3-4-5- ?4 is
evident in the pes but no skeletal pattern is discernible in the
hoof-like manus.

In preparing the skeletal restorations of C. parvum and C.
eyermani I have indicated joints at localized thickenings in the
sole wherever possible, and have extrapolated as little as pos-
sible from the structure of other species. In C. parvum a slight
differentiation in the coalesced pad of pes digit V suggests the
presence of three phalanges, so by analogy the same number is
hypothesized for C. eyermani.

Classification of Chirotheriids

Peabody, working chiefly with Lower Triassic species, has sep-
arated the better-known chirotheriids into a typical or large-
manus group and a specialized small-manus group. Ratios of
manus to pes area in representative members of each group are

482 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

tabulated below, in order of decreasing relative manus size.
(Areas include only the digits and metapodio-phalangeal pads.
These ratios, based on single manus-pes sets, are not necessarily
typical of the species listed.)

Large-Manus Small-Manus

C. lulli Bock 1 :2 C. coltoni Peabody 1 :5.7

C. storetonense Morton C. marskolli Peabody 1 :5.7

ca. 1 :3 C. lomasi Baird 1 :6.6

C. minus Sickler 1 :3

C. wondrai Heller 1 :3.4:

C. harthii Kaup 1 :3.6

In Chirotherium parvum the type manus is too incomplete for
areal measurement but ANS 15210 furnishes reliable figures of
25 cm. 2 for the manus and 112 cm.^ for the pes. (Bock's meas-
urement of 85 cm.- for the pes apparently omits the "thumb"
pad.) The resulting manus-pes ratio of 1:4.6 falls between the
ranges of the two groups. Except for the closeness of digit V
to its fellows the manus of C. parvum is more similar to that of
the large-manus species than to the compact, hoof-like forefoot
of the small-manus group. Unfortunately, nothing is known of
the manus in C. eyermani.

The shortening of pes digit IV, moderate in C. eyermani but
extreme in C. parvum, would seem on typological grounds to ally
these species with the small-manus group, in which such a
shortening is characteristic. In terms of function, however, this
condition is obviously essential to the maintenance of symmetry
in any walking foot in which the fifth digit has undergone
reduction. Other examples are found in the feet of dinosaurs,
crocodiles and birds (where apparent exceptions such as Gavia,
Pelecanus and Hesperornis prove the rule, since walking is not
the primary function of the foot). Thus the shortened fourth
digit in the Milford species is functionally correlated with the
vestigial fifth digit and indicates parallelism rather than affinity
with the small-manus group.

The Level B chirotheriids thus appear to represent a lineage
distinct from the small-manus group although paralleling it in
adaptive modifications and similarly derived from a primitive
large-manus stock. This lineage has been progressively modified

TRIASSIC REPTILE FOOTPRINTS 483

to a point where it must be considered a distinct sub-generic
group, characterized by the short, oval, heel-like fifth pes digit.

This group, here recognized in North America for the first
time, is represented by specimens from several horizons in the
Middle Keuper of Germany (Heller, 1952). The oldest of
these, an imperfect pes imprint from the Blasensandstein of
Franken, was described by 0. Kuhn (1936) as Chirotherium
sp. In size and proportions it shows a decided resemblance to
the Level B species, particularly C. eyermani, differing chiefly
in the greater length of digit V.

A rather similar but slightly younger form is Chirotherium
thiiringiacuni Rlihle von Lilienstern (1938) from the Platten-
sandstein (km 4 zeta) of Thiiringen. This species recalls C.
eyermani in its short, divergent pes digit I-IV group. It re-
sembles C. parvum in its round, apparently clawless digit tips;
and the manus with its plump, clawless toes and close-set, medi-
ally shifted digit V is strikingly similar.

Slightly higher in the Middle Keuper, in the Semionoten-
Sandstein of Franken, occurs a group of chirotherioid footprints
which Beurlen (1950) has named Brachychirotherium hassfur-
tense. Though the specimens figured are somewhat heterogeneous
they all show a foot structure like that described above, with
apparently clawless digits and an abbreviated "thumb." The
latter appears so inconsistent in its size, position, and orienta-
tion, and the relative lengths of the other digits are so variable
from specimen to specimen, that definitive diagnosis of the
species (singular or plural) is impossible. Unfortunately, all
these German forms have been described from individual foot-
prints rather than analyzed on the basis of a composite of each
population. Specific distinctions are by no means clear, particu-
larly as no types have been designated, and the differences care-
fully tabulated by Heller may or may not be significant. Never-
theless these Middle Keuper species from Germany are clearly
referable to the same group as C. parvum and C. eyermani.

For this group Beurlen has established the separate genus
Brachychirotherium. Since the differences between this and the
large-manus group are little greater than those which distinguish
the small-manus group from both, I prefer to broaden slightly
the scope of the form-genus Chirotherium to include a third

484 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY

sub-generic group for which Beurlen's name may appropriately
be retained. Following the example of Peabody I assign this
group no formal Linnaean status.

Evolutionary Tendencies

The modifications seen in this brachychirotherian group are
clearly consistent with the general evolutionary trend of Keuper
chirotheriids which I have previously outlined (1954, p. 188) :
the tendency of an osteologically pentadactyl stock to become
functionally tetradactyl through elimination of the lateral prop-
ping function of the fifth pes digit. In the brachychirotherian
group this was being accomplished by several means : by shifting
the distal end of the unshortened metatarsal V more or less
medially, beneath and behind the shaft of metatarsal IV; by
progressively shortening and straightening the fifth digit to
minimize its lateral projection; and ultimately by incorporating
the metatarsal and phalanges into a single, inflexible heel
structure. During this process a gradual shortening of digit
IV tended to maintain the symmetry of the foot. Concomitantly
the manus became smaller and more compact.

Although — most unfortunately — we have no trackways from
which to determine the locomotor habits of the group, it seems
inescapable that these modifications promoted more efficient
locomotion, presumably cursorial and (as Riihle von Lilienstern
points out) showing bipedal tendencies in some members. In-
creasing competition from the expanding and diversifying stocks
of saurischian dinosaurs may well have channeled chirotherian
evolution along these lines.

Changes in foot structure and locomotor pattern are of course
only a part of the overall adaptive modification which must have
taken place in the brachychirotherian group. Only a part, yes :
but surely an important one, for, as I have shown (1954, p.
174), convergent modifications were independently taking place
in Keuper representatives of both the large-manus and small-
manus groups. Apparently the chirotheriid locomotor equipment
of early Triassic time had been found inadequate for the con-
ditions of mid-Keuper life.

The chirotheriid response to changing conditions was, how-
ever, insufficient. We have noted that even in the most special-

TRIASSIC REPTILE FOOTPRINTS 48;")

ized species, C. parviim and C. eyermani, the once thumb-like
fifth digit was still functional in modified form. Dinosaurs and
small pseudosuehian or proto-erocodilian types such as Batracho-
pus, in which locomotion had become more efficient through the
loss or atrophy of the fifth digit, eventually replaced Chirother-
iinii. Huge size and bipedality (along with other specializations
yet unknown) permitted a lone descendant of the chirotheriid
stock, Ofozonni, to survive until the close of Newark time — but
that is another story.

Osteological Correlations

That chirotheriid trackways are the products of various
pseudosuehian reptiles is now" generally accepted. Beyond this
premise the foregoing discussion has avoided assumptions as
to the identity of the Milford trackmakers, attempting instead
an objective analysis of the ichnological evidence as such. The
possible position of these trackmakers among the Pseudosuchia
can now^ be examined.

At present the pedal skeletons of most Upper Triassic pseudo-
suchians are too inadequately known to permit useful compari-
sons with the three Milford species of Chirotheriitm. The rela-
tively conservative large-manus species C. lulli has been inter-
preted as a small, persistently quadrupedal ornithosuchid not
too different from the presumably ancestral form Eupar'keria;
but closer comparisons are impossible. Among middle to late
Triassic reptiles which might be compared to the brachychiro-
therian species only two, Prestosuchus and Typotliorax, are
represented by even relatively complete foot material.

As restored by Peabody (1948, fig. 39 B-C^) the pes of Presto-
suchus is broad and short-toed, suggesting (as Peabody notes)
the brachychirotherian C. thuringiacum Riihle. Lack of the tip
of digit IV and all phalanges of digit V, however, precludes for
the present any correlation of Prestosuchus w^ith one or another
of the chirotheriids. Certainly the phalanges are relatively much
shorter than those restored for the Milford footprints. As Huene
(1944) records only a single phalanx from the manus, Pea-

3 Peabody has followed Huene in reversing the proximal overlap of the meta-
tarsals although the natural relationship is preserved in the specimen itself ^ —
cf. Huene, 1944, pi. 20, hg. 4 a-b.

486 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

body's very plausible reconstruction of the manus digits cannot
be used for comparison.

Despite the strong contrast in habitus between the relatively
narrow-bodied, lightly-armored, predaceous Prestosnchus and
the broad-backed, heavily-armored, necrophagous Typothorax
(Sawin 1947), the two are striking^ similar in phalangeal
pattern. Thus the general aspect of Typothorax is brachychiro-
therian-like but its short phalanges seem to eliminate it from
comparison with the Milford footprints. The manus is decidedly
chirotherian in form although its fifth digit appears to have
been somewhat weaker than that of most chirotheriids. Areas of
the digit I-IV groups in manus and pes bear a ratio of about
1 to 3, a ratio found in C. thuringiacum and some large-manus
chirotheriids. If brachychirotherian trackways were available
the pedal similarities between Prestosuchus and Typothorax
should cause little difficulty in correlation, for the swift-striding
predator must have left a much narrower trackway than its
broad-beamed, ponderous relative. But pending the discovery of
such additional evidence the systematic positions of the Level
B pseudosuchians must remain uncertain.

Order THECODONTIA

Suborder PHYTOSAURIA

Form-family APATOPODIDAE Baird, nov.

Genus APATOPUS Baird, nov.

Type speeioM A. Imeatus (Bock), 19."2; family and genus monotypie.

Diagnosis. Quadrupedal trackway wdth pace angulation about
108°, pes but not manus toed-out. Feet pentadactyl with well-
developed articular swellings and sharp claws ; manus short and
symmetrically radiate, pes long and narrow^ with digits in order
of increasing length V-I-II-III-IV.

ApATOPUS LINEATUS (Bock)

Figures 7, 8 ; Plates 3, 4

Otosoum (?; lineatus Bock, 1952, pp. 408-409, pis. 48, fig. 1 and 46, fig. 1
(niistitled "Type No. 15230" for LC S489). Probable earlier syn-

TRIASSIC REPTILE FOOTPRINTS 487

onyniy in Table 1 ; the variety of past assignments suggested the
generic name. Diagnostic characters are those of the genus.

Type. LC S490. a left maniis-pes set (Bock's type) plus MCZ
212, the succeeding right set of the same trackway.

Hypodigm. The type, S489, and MCZ 213, presumably three
adults ; AMNH 1789 and 2258, smaller individuals ; AMNH 2259
and MCZ 214, juvenile (same individual?). Source data as for
A nch isa u rip u s ni ilfo rclensis.

Trackway. By a remarkable and fortunate coincidence a tiny
contact area between corners of the Lafayette and Harvard
slabs permits the reconstruction of the type trackway. (This
contact crosses the "thumb" of the Chirotherium parvum type,
cf. Figure 5 B.) Trackway parameters determined by direct
measurement or by extrapolation (*) are tabulated below. In
the first three the reference point is the center of metatarso-
phalangeal pad III :

Pace, angular 42.5 cm.

Stride *70 cm.

Pace angulation *108°

Gleno-acetabular length (approximate) *52 cm.

Interpes width (between median margins) 13.7 cm.

Pedal morphology. The additional material now available
permits analysis which was impossible with the incomplete, ob-
scure, single manus-pes set on which the species was based. A
composite restoration derived from all this material is presented
in Figure 7. The digits of both manus and pes are shod with
domed, oval articular pads separated by slender waists, so that
joint positions are apparent in profile as well as in section. The
claws are slender and rather crocodilian in the manus but appear
heavier and more triangular in the pes, where rotation during
withdrawal has obscured their exact form. As the plates show,
laterally flexible ungual articulations permitted much lateral
movement in both manus and pes claws.

The manus is nearly symmetrical around digit III, the meta-
carpo-phalangeal pads forming a regular semicircle, so that in
form and proportions it recalls the manus of Mesosaurus —
an aquatic form. A phalangeal formula of 2-3-4- ?5-?3 is indi-
cated. In normal function the manus was digitigrade and only
in the deepest imprint did the palm register (Plate 4, figure 1).

The pes with its long fourth digit and peculiar sole is quite

488 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

A

CM.-r

5-

10-

15-^

Fig. 7. Apatopus lineatus (Bock), composite restoration based on type
(S490 + MCZ 212) with parts of digits IV and V restored from other
specimens, x 1/2. Arrow represents midline of trackway.

TRIASSIC REPTILE FOOTPRINTS 489

distinct from other Triassic footprint types. At first glance it
appears to show affinities with primitive reptilian pedes of the
pelycosaur or captorhinomorph type, but the fact that meta-
tarsal IV is slightly shorter than III (rather than the reverse)
indicates that digit IV has been secondarily lengthened by hyper-
trophy of its phalanges. A similar secondary lengthening of the
phalanges of digit IV occurs in the manus of the marine turtle
Chelonia mydns (cf. Abel, 1912, fig. 111). Aquatic adaptation
is the most obvious explanation for such a modification, and this
interpretation is supported by the evidence of interdigital webs.
Although the length and position of the short fifth digit are evi-
dent its structural details are obscure ; my conjectural restora-
tion shows four phalanges, making the pes formula 2-3-4-5- ?4.

The extent of interdigital webbing in Apatopus is uncertain.
])articularly because the pes in walking was often laterally com-
pressed so that web margins would have left no mark. The best-
represented web connects the bases of pes claws III and IV, an
arrangement quite in contrast to that in living crocodilians, in
which the deeply recessed webs occupy only part of the inter-
digital area. Additional (though circumstantial) evidence for
the presence of webbing is the fact that digits IV and V are
adequately recorded only in the deepest footprint, which sug-
gests that distribution of the animal's weight over a w^ebbed
area effectively reduced the depth of impression. Here as in
other fossil footprints, unfortunately, the evidence is less than
satisfactory.

Traces of the squamation are preserved only in the juvenile
MCZ 214 (Plate 4, figure 1). Here the digits of manus and pes
show a fine beaded or pustulose texture which compares closely
with the skin of corresponding areas in a juvenile Gavialis
gangeticus.

Except in the deepest impressions the sole is very indistinctly
recorded. Some understanding of it can nevertheless be obtained
by collating outlines printed from latex molds of several speci-
mens. On the medial margin a bulge marks the position of the
first metatarsal's proximal head, and opposite this is a circular
pad which must have underlain the fifth metatarsal. On this
basis the first four metatarsals have been restored in normal
alignment. Its distinct individual pad suggests that the proximal
end of metatarsal V was less elevated than that of metatarsal IV ;

490 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

if SO, then the fifth metatarsal probably articulated with the
tarsus by a hooked, dorso-medially directed process. Postero-
medial to this area a large, subcircular calcanear pad forms
the heel of the footprint. The possible presence of a tuber eal-
canei is suggested by the relatively deep impression of this pad.
Though plausible, this interpretation of the tarso-metatarsal
area rests on admittedly tenuous evidence.

Variation. Aside from striking variations in the manner of
impression, which are apparent in the plates, the Apatopxis
footprints reveal only minor differences among the six (or so)
individuals of the population. Chief of these is size : the small-
est individual, AMNH 2259, has a pes length just half that of
the type (as restored). Osteological analysis reveals the struc-
tural unity of these footprints which might otherwise be as-
signed to a number of form-species.

Affinities of Apatopus

Examination of Triassic footprints from every continent
yields nothing similar enough to Apatopus to justify comparison
in terms of footprint taxonomy. We may more profitably in-
vestigate the nature and systematic position of the trackmaker.
In its foot structure and body proportions Apatopus shows closest
affinities to quadrupedal members of the Subclass Archosauria —
i.e. the Phytosauria, Pseudosuchia and Crocodilia. These groups
will be considered in reverse sequence, beginning with the best
known.

Locomotion in crocodiles, which has been investigated by Abel
(1912, p. 217), Huene (1913), and Schaeffer (1941, p. 443 ff.),
is of three types: a slow, lizard-like dragging of the body, a
spring in which all four feet thrust simultaneously, and a strid-
ing gait in which the belly is well above the ground and only
the tail-tip leaves a mark. Of these gaits only the last would
produce trackways comparable to that of Apatopus; thus neither
the "Gehspur" nor the "Laufspur" which Huene figures is
pertinent to this discussion. An example of the true striding
gait in a juvenile Alligator mississipiensis has been recorded
on film by Schaeffer. The resulting trackway shows a pace angu-
lation of only 80°, i.e. the stride is little longer than the pace. In
evaluating this record the smallness of the alligator and the

TRIASSIC REPTILE FOOTPRINTS

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492 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

fact that it was walking on glass must be considered. For ex-
ample, the 20° to 30° outward rotation of the pes between
implantation and withdrawal can hardly be normal : no such
rotation is evident in the alligator trackway figured by Jaeger
(1948, pi. 170 c), and only in very muddy footing did the pes
of Apatopus rotate a few degrees outward (cf. Plate 4, figure
2). I believe that on a fairly firm surface a crocodilian in rapid
motion would produce a trackway rather similar in proportions
to that of Apatopus. We have already noted the crocodilian
habitus indicated by the aquatic adaptations in the feet of the
Milford traclanaker.

Despite, these similarities in habitus Apatopus can hardly
have been a crocodilian by heritage. The ancestral crocodilian
Protosuchus from Upper Triassic or Lower Jurassic beds of Ari-
zona had already developed an essentially modern type of foot
with an elongate metatarsus, reduced fourth and rudimentary
fifth digits, and a formula of 2-3-4-4-0 (Colbert and Mook, 1951).
These characteristics were probably inherited without much
modification from pseudosuchian ancestors and have been trans-
mitted without much modification to the living crocodilian.s.
Even the aberrant pes structure in the marine thalattosuchians
can be more readily derived from the Protosuchus pattern than
could that of Apatopus. Another point of distinction is the out-
turned manus which characterizes crocodilian trackways. Thus
the Milford trackmaker cannot be referred to any crocodilian
lineage.

Among trackways which have been ascribed to pseudosuchians
certain members of the Batrachopns group approach the Apato-
pus pattern. Nevertheless the reconstructed feet of these forms
— and the few known pseudosuchian foot skeletons — are dis-
tinctly different, particularly in the fourth and fifth pes digits.

In the Phytosauria we find the logical correlative for a
trackmaker which is crocodilian in body form and presumed
habitus but not in skeletal structure. The comparative propor-
tions of the crocodilians Gavialis, Tomistoma and Alligator, and
the phytosaurs Eutiodon, Macliaeroprosopus and Mystriosuchus
have been analyzed by Colbert (1947), who attests to the re-
markable parallelism noted by earlier authors. Very little articu-
lated phytosaurian foot material, unfortunately, is available for
comparison with the footprints. The manus is best represented

TRIASSIC REPTILE FOOTPRINTS 493

in Mystriosuchus (McGregor, 1!)06, fig. 10), in which — like
Apatopus — the metacarpal bundle is nearly bilaterally sym-
metrical and digit V bears three phalanges which are abont
twice as long as they are wide. Further comparisons cannot be
made. McGregor's reconstruction of the pes in Ridiodon, based
on incomplete, scattered material of several individuals, is
remarkably similar to my restoration of Apatopus — which (ex-
cept where drawn in dashed lines) was quite independently
derived from the footprints alone. If Apatopus is indeed a
phytosaur, then Dr. McGregor proves to have been indeed a
prophet worthy of honor.

Correlation with the Phytosauria is supported by all the
evidence and contradicted by none; it may therefore be taken
as a working assumption. Possible equivalents exist in the two
pln-tosaurs of the Newark series, Eutiodon and Clepsysaurus
{'i=Machaeroprosopus) . The significant differences in skeletal
proportions between these genera would presumably be reflected
in their trackways and may someday permit positive assignment
of Apatopus lineatus to one or the other. Present knowledge,
however, does not justify comparison of the single, partial
Apatopus trackway with the composite skeleton of Rutiodon
from North Carolina (Colbert, 1947) as opposed to incomplete
skeletons from Arizona which may be congeneric with Clepsysau-
rus (Camp, 1930). Geographic distribution of the Newark
phytosaurs offers a possible guide to the identity of A. lineatus,
for Colbert and Chaffee (1941) concur with Camp's conclusion
that all the phytosaur remains from Pennsylvania, New Jersey,
and Connecticut represent Clepsysaurus while the North Caro-
lina finds all pertain to Rutiodon.^ Apatopus lineatus would
thus appear to be a correlative of Clepsysaurus.

So far as I know these Milford specimens are the only phyto-
saur footprints yet described.^ Paradoxically, phytosaur bones
are the commonest skeletal remains of both the Chinle formation
of the Southwest and the Newark series of the East (excluding.

6 H. E. Wanner's (1926) specimens of "Rutiodon caroUnensis" from the Newark
series of Yorlj County, Pennsylvania — overloolied by the above-mentioned authors
and by the Hay-Camp bibliographies — merit further study.

7 An isolated pes imprint, considered by Bock (19o2, pi. 46, fig. 2) to be
probably that of a large parasuchian of the Clepsysaurus type, bears no resem-
blance to Apatopus. So far as its preservation permits comparisons, it seems
more similar to ornithoid footprints of the Plectropterna type from the Con-
necticut Valley.

494 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

significantly, the Meriden and Portland formations). In the
ecology lies the probable explanation for this discrepancy :
Triassic footprint assemblages as a rule represent terrestrial
faunas while the fossil deposits contain a disproportionate num-
ber of aquatic forms. Thus the presence of several phytosaurs
in a dinosaur-pseudosuchian-lacertoid footprint faunule at Mil-
ford is rather anomalous.

REPTILIA LACERTOIDEA INCERTAE SEDIS

Genus RhYNCHOSAUROIDES Beasley w Maidwell, 1911

Fontopus Nopcsa, 1923, p. 141. Keiiper of England.

Akropus Riihle von Lilicnstern, 1939, p. 298. Bunter of Germany; Moenkopi
of Arizona (Peabody, 1948, p. 319 ff.).

Hamatopus Riihle von Lilienstern, 1939, p. 319. Bunter of Germany.

TJiyncliocephalichnus Huene, 1941, p. 14. Keiiper (Verrucano) of Italy;
Keuper of Germany (Heller, 1956).

Eurichnus Lull, 1942A, p. 502; Branson, 1947, p. 590. Lower Triassie
(Chugwater) of Wyoming.

The type species is here designated as R. rectipes Beasley in Maidvrell, the
type specimen of which is a manus-pes set from the Keuper of Dares-
bury, England (Dr. Ricketts' collection, University of Liverpool
Geology Department).

RHYNCHOSAUROIDES HYPERBATES Baird, n. Sp.

Figures 9 A, 10

Type. ANS 15210, a trackway of one pes and three manus
imprints associated with the type of Chirotherium "copei."

Hypodigm. The type ; an isolated left pes on the same slab ;
MCZ 210, a partial left pes associated with C. parvum.

Diagnosis. Manus i}road with little disparity in digit lengths :
gait digitigrade with hallux non-functional and proximal ends
of proximal phalanges normally carried clear of the ground.
The specific name alludes to the overstepping of manus by pes.

Morphology. The trackway is rather narrow, showing a stride
of 44 cm., forelimb paces of 23 and 25.5 cm., and a pace angula-
tion of 130° between manus imprints. (In these measurements a
3-cm. fissure filling which crosses the trackway is ignored, as
the relative ages of trackway and fissure are unknown.) As in

TBIASSIC REPTILE FOOTPRINTS

495

Fig. 9. A. Bhynehosauroides hyperbates n. sp., composite restoration
of manus (at left) and pes in normal position. B-C. Small Bhynehosauroides
footprints from Milford: B, (ANS 15210); C. manus imprint (MCZ 213).
D. Manus imprint from Trostle quarry near York Springs, Adams Co., Pa.
(Carnegie Museum 12082). E. Manus of Monjurosuchus reconstructed from
Endo's photograph. F. Manus and pes of Polysphenodon miilleri Jaekel,
modified from Huene. All x 4/5.

496

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

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TRIASSIC REPTILE FOOTPRINTS 497

Other Triassic lacertoid trackways the pes is strongly out-turned
and evidently overstepped the manus as it implanted.^ The
trackmaker, which is restored in walking pose in Figure 10, had
a gleno-acetabnlar length of about 18 cm. and a total length of
perhaps 75 cm.

The foot structure, revealed by comparing outlines printed
from latex molds of all the imprints, is decidedly lacertoid.
Well-developed articular swellings indicate an apparent pha-
langeal formula of 1-2-3-4- ? in the manus and ?-2-3-4-? in the
pes. This formula must be incomplete, for the obvious reptilian
character of the feet connotes a normal formula of 2-3-4-5-3(4).
Like the structurally similar Moenkopi genus Rotodactylus (Pea-
body, 1948, p. 329), the Milford form must have been digitigrade
to such an extent that each proximal phalanx was elevated with
its metapodial ; normalh^ only the padded distal articulation of
phalanx 1 touched the ground. All the digits recorded bear
sharp claws, either extended or curved toward the trackway
midline.

The pollex is represented only by claw impressions. Nothing
which can be identified as the imprint of a fifth manus digit
is visible. The absence of any hallux imprint and the disparity in
length of the pes digits indicate that the hallux was non-func-
tional. Pes digit V has left no unequivocal record, but obscure
marks on MCZ 210 and ANS 15210 may have been made by a
lacertoid, laterally projecting minimus. Both these marks are
shown, for what they may be worth, in Figure 9 A.

Rhynchosauroides sp.
Figure 9 B, C ; Plate 3, Figure 2

A pair of minute footprints on the ANS slab, unfortunately
too incomplete for reliable analysis, and indistinct trackways of
comparable size on a slab with Apatopus lineatus (MCZ 213)
prove the presence of another species of Rhynchosauroides in the
Level B faunule. Smaller size and greater disparity in the
lengths of the manus digits distinguish it from B. hyperhates.

8 1 have seen no evidence in Pennsylvanian, Permian, or Triassic lacertoid
trackways that the pes rotated nearly 90° outward during the propulsive phase
as Schaeffer (1941, p. 440) has observed in experiments with Anolis. Except
tor an occasional rotational slippage of a few degrees in unusually soft mud
the pes remained in implantation position until withdrawn.

498 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Relationships

Lacertoid trackways of this sort are familiar companions of
Chiroiherium in the Lower and Upper Triassic of both Europe
and America. As indicated in the synonymy, a number of generic
names have been applied to them. The differences between these
nominal genera are, however, slight : for example, Riihle von
Lilienstern distinguished Ilamatopus from Akropus largely by
the more hooked appearance of its digit tips. A more practical
treatment here proposed is to consider these forms as species of
a single form-genus Rhyncliosaur aides which is comparable to
Chirotherium in its range of variation. This form-genus, of cos-
mopolitan distribution in continental Triassic beds, undoubtedly
represents several reptilian genera; but until more is known of
Triassic lacertoid feet and footprints there seems little advantage
in proliferating form-genera on differences of uncertain diag-
nostic value.

Our lack of data on the fifth digits of manus and pes hinders
comparison of Ehynchosauroides hyfjerhatcs to other species. Its
closest similarities, however, are to the footprints from the
English Keuper which were described by Beasley (1905) and
comprehensively reviewed by Maidwell (1911, 1914). The
smaller Milford lacertoid is less similar to the larger one than to
Rhynchosauroides [Rhynchocejyhalichnus] franconicus (Heller,
1956; see also Haarlander, 1938) from the Keuper of Germany.

Such trackways have never been found in the well-known
footprint faunas of Portland and Meriden age in the Connecticut
valley, though they are not uncommon in the Brunswick and
Lockatong formations of the Pennsylvania-New Jersey basin.
The Milford forms cannot adequately be compared with Rhyn-
chosauroides ['^ Orthodactylus"Y howelli (Bock) from an earlier
horizon of the Brunswick near Schwenkville, Pennsylvania, until
better-preserved material is available. An isolated Rhyncho-

9 This form has nothing to do with Hitchcock's Orthodactylus from the Port-
land formation of Gill, Massachusetts. The type specimen of the type species
O. florij^rus is an obscurely recorded trackway of pseudosuehian or proto-
crocodilian tvpe, having the manus anterior to the pes and strongly out-turned as
in the contemporary (Tit HOi/ieroide«. _ Superficial similarity of the digital gouge-
ma " ■ - - -"

dactylh.. -_ — . - .„,,-.

and equally indeterminate were made the type of a new species, Procolophoinpus
conhuenti 'Bock (p. 419), ascribed to a plump, slow-moving reptile which jumped
quadrupedally without leaving manus imprints.

TRIASSIC REPTILE FOOTPRINTS 499

sauroides maniis imprint of Brunswick age from the Trostle stone
quarry near York Springs, Adams County, Pennsylvania, is
shown in Figure 9 D ; and another of Lockatong age from
Phoenixville, Pennsylvania, has been figured without name ])y
Bock (1952, pi. 48, fig. 2). Aside from some undescribed track-
ways from the Doclmm group of the Cimarron Valley in north-
eastern New Mexico (AMNH 6338), these are the only records of
Rhynchosauroides in the Upper Triassic of North America.

The systematic position of the trackmakers is debatable. Foot-
prints of this sort have been referred by various authors to the
Protorosauria, Eosuchia, Pseudosuchia, Rhynchocephalia and
Lacertilia: and indeed each of these groups has Triassic repre-
sentatives which would have made lacertoid footprints. Among
the Protorosauria the tanj'stropheid Macrocnemus from the Al-
pine Middle Triassic (Pe^^er, 1937) is similar in size and pro-
portions. Its reduced fifth digits of manus and pes exclude it
from correlation with the European species of Rhynchosauroides,
and its greater disparity in the lengths of the manus digits
distinguishes it from the larger Milford form. A similar dis-
parity characterizes the manus of Trilophosauriis from the
Upper Triassic (Dockum) of Texas (Gregory, 1945) ; but its pes
digits are much less unequal in length. The proximal end of
ungual II lies opposite those of phalanges III-3 and IV-2, while
in the footprints it lies opposite the proximal end of III-2 and
the waist of lV-1, much as in the Permian Protorosaurus. But as
the known Triassic protorosaurian genera undoubtedly consti-
tute only a fraction of the number that once lived, the group
cannot be disregarded as a possible source of Keuper lacertoid
trackways.

Too little is known of limb structure in the Eosuchia to permit
useful comparison.

Peabody (1948, p. 337) has referred his Lower Triassic foot-
print genus Rotodactylus to the Pseudosuchia on the basis of
its long-striding gait with pendulum limb movement, its semi-
bipedal concentration of the body weight on the long hind limbs,
and its digitigrade pes with posteriorly rotated, prop-like fifth
digit. Isolated mauus-pes sets of Rotodactylus appear decep-
tively lacertoid (especially where the fifth digit has failed to
impress), but the whole organization of the animal is significantly

500 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

different from that of lizard-like reptiles/" Aside from its over-
stepping gait and similar digitigrade specialization the Milford
trackmaker appears to be basically unlike Rotodactylus. No
comparable pseudosuchian skeleton is known.

The case for the Rhynchocephalia as a source for Triassic
lacertoid footprints, reviewed by Huene (1938, 1941) and Pea-
body (1948), dates historically from Richard Owen, who in
1842 assigned the small footprints of the Grinsill quarries near
Shrewsbury to Bhynchosaurus articeps on the grounds that the
bones and footprints correspond in size and occur in the same
beds, to the exclusion of other bones and footprints. This correla
tion was accepted by subsequent students of British Keuper foot-
prints and indeed motivated Beasley to coin the name Bhyncho-
saur aides.

A basic similarity certainly exists between Rhynchosauroides
footprints and the feet of Bhynchosaurus as restored by Smith
Woodward (1907). But closer inspection shows that nearly all
the footprints were made by long-legged, slender-toed reptiles,
whereas Bhynchosaurus is comparatively short-legged and broad-
toed as a modification for aquatic life. At this point it may be
illuminating to note that in the same year in which he proposed
the Bhynchosaurus correlation Owen was asserting, on essentially
similar grounds, that Chirotheriuni footprints represented Laby-
rinthodon. The great contrast (now understood) between the
feeble-limbed, bottom-crawling " Lahyrinthodon" and the up-
right-limbed, semi-cursorial Chirotheriuni points up the ecologi-
cal unsoundness of both correlations: the genera common in the
Keuper bonebeds represent an aquatic facies-fauna while the
footprints record a terrestrial facies-fauna.

Among terrestrial rhynchocephalians we may eliminate the
Rhynchosauridae because of their un-lacertoid proportions and
short, hea\y phalanges. The Sphenodontidae, however, offer
closer comparisons. Proportions similar to those of the Milford
trackmaker are found in the Upper Jurassic Homoeosaurus; the
Manchurian sphenodontid Monjurosuchus (Endo, 1940) is rather
similar in manus structure (Figure 9 E) although its body pro-
portions suggest that the pes probably did not overstep the

10 Bock's attempt (1952, p. 422) to synonymize Rotodactylus with the
araeosceloid Lower Permian footprint •Ichnium gampsodactijhnn" — more cor-
rectly termed Dromopus [Saurichnites] lacertoides (Geiuitz), n. comb. — over-
values the apparent similarities at the expense of fundamental ditTorences in foot
structure and tracljway pattern.

TRIASSIC REPTILE FOOTPRINTS 501

manus as much as in Rhynehosanroidcs hyperhates. Upper Tri-
assic genera are (at this writing) less completely known, but
the limbs of Polysphenodon compare favorably (as Huene points
out) with footprints of the Rhynchosauroides type from the
Keuper of Italy. As shown in Figure 9 F, the phalangeal pattern
of Polysphenodon also corresponds approximately to that of the
larger Milford lacertoid. The publication of current research
on several skeletons of Glevosaurus, a sphenodontid from Keuper
fissure fillings in Gloucestershire (Robinson, 1955), may well
provide the means to a better understanding of Rhynchosau-
roides.

Triassic Lacertilia, a group in which major advances in knowl-
edge are in progress, may similarly prove to correlate with
some of the lacertoid trackways which I have brigaded under the
name Rhynchosau7'oides. On the evidence now available R.
hyperhates is assigned tentatively to the sphenodontid Rhyncho-
cephalia, but laeertilian origin remains a distinct possibility. At
present the various trackways cited above constitute the only
evidence of rhynchocephalians or lacertilians in the North Amer-
ican Triassic.

CORRELATIVE AGE OF THE MILFORD FAUNULES

Research on several problems of Pennsylvanian and Permian
ichnology, still unpublished, indicates that fossil footprints —
especially footprint faunas — can be used in a limited way for
stratigraphic correlation. As footprint species can at best be
equated only with generic units of tetrapods they cannot ap-
proach the diagnostic precision of skeletal remains. Assemblages
of footprints are of course subject to the same ecological bias
as assemblages of skeletons, and may be equally deceptive in
suggesting the non-contemporaneity of beds which are actually
equivalent in age.

Despite such limitations ichnological evidence can be a valid
basis for correlation; indeed, it is sometimes the only basis
available. One pertinent example is the presence of similar
Chirotherium harthii faunas in the Bunter of Germany and the
Moenkopi of Arizona, which (with other evidence) .establishes
the equivalence of these strata. Consequently when a specimen
of C. barthii which is indistinguishable from the Arizona ma-

502 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

terial appears (under the name ''C. higuerensis") in so-called
Upper Permian beds of Argentina (Rusconi, 1951) we need feel
no doubt that the beds are in fact Triassie/^

The amount of material now available and the size and diver-
sity of the combined Milford faunules justifies an attempt at
correlating them with faunas of other areas. In this assay foot-
prints will be compared with footprints and bones with bones :
for in the past, uncritical herpetological interpretations of mis-
identified and uuanalyzed footprints have too often proved
misleading in stratigraphic work.

Two views on the correlative age of the Milford footprints
have been advanced : one, that the footprint assemblages of Mil-
ford and Whitehall, New Jersey, York Springs, Pennsylvania,
and Aldie, Virginia, compare closely with the youngest foot-
print fauna of the Portland formation in Massachusetts (Bock,
1952, pp. 429-430) ; the other, that the Milford faunules correlate
neither with the Portland nor the underlying Meriden but with
the New Haven, lowermost of the Connecticut Valley formations
(Baird, 1954, pp. 184-187). Let us examine these faunules, be-
ginning with the southernmost.

The only specimen from Aldie available to me (MCZ 236) is
a well-preserved pes imprint, 35 cm. long including the hallux,
of Anchisauripus tuherosiis (Hitchcock). This species ranges
from the lower Meriden to the upper Portland formation. Of the
Aldie fauna Gilmore (1924) says:

"Three-toed imprints predominate though they vary in size
from a length of three to fourteen inches. A few tracks were
noticed having four toes, evidently terminated with wide, flat
unguals. All of these are probably of dinosaurian origin, but
a few small 4- or 5-toed tracks with traces of sharp claws per-
haps pertain to some other group. ' '
Until this important assemblage has been thoroughly studied
no valid comparisons can be made.

Specimens from the Gettysburg shale of the Trostle stone
quarry near York Springs, Adams County, Pennsylvania, are
figured by Stose and Jonas (1939, pi. 22) ; others are preserved
at the Carnegie Museum. In addition to invertebrate trackways

11 Siuce the foregoing was written Peabody (19.55) has published the same
observation and noted the importance of this evidence on Triassic faunal distribu-
tion.

TRIASSIC REPTILE FOOTPRINTS 503

and some indeterminate vertebrate tracks the faunule contains
three genera: (1) a uniquely quadrupedal dinosaurian type re-
sembling Anchisauripus gwyneddensis Bock from the Lockatong
formation, (2) a Blujnchosauroides (Figure 9 D) comparable to
those from the Brunswick of Schwenkville and Milford and the
Lockatong of Phoenixville, and (3) large footprints consisting
of a sub-circular plantar pad close behind an arc of broad ungual
depressions, rather suggestive of a dicynodont foot. Nothing like
any of these forms occurs in the Connecticut Valley.

The Whitehall footprints are known to me only from the
thoroughly unreliable faunal list of C. H. Hitchcock (1889).
Further discussion is useless until the material can be restudied
by modern methods.

Only one of the Milford species, Anchisauripus parallelus, also
occurs in the Connecticut Valley. The other dinosaurian tracks
are distinctly different from those of the Meriden and Portland
formations. Chirotherium footprints, which at Milford record
three genera of pseudosuchian reptiles, are unknown in New
England ; so are trackways of the Apatopus and Rhyrichosau-
roides types. Their absence there is real and cannot be ex-
])laiued as the result of inadequate collecting or superficial study
— far from it 1 In the absence of any evidence of barriers be-
tween the New Jersey-Pennsylvania basin and the ancient Con-
necticut Valley trough, the conclusion presented in m.y previous
j)aper seems inescapable : the footprint beds of the upper Bruns-
wick formation at Milford, New Jersey, antedate the Portland
and Meriden formations and more probably correlate with the
New Haven arkose of Connecticut. The most comparable foot-
prints in the European sequence occur in formations of the mid-
dle to upper Middle Keuper of Germany.

REPTILES OF THE BRUNSWICK FORMATION

To recapitulate, the Milford footprints comprise three mu-
tually exclusive faunules from three horizons of the Brunswick
formation. What these footprints signify in herpetological terms
is summarized, so far as known, in Table 3. These footprints
probably represent only a few of the different types of reptiles
which inhabited the area, and the reptiles themselves only a
.segment of the biota. Nevertheless a partial account of the

504

BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

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TRIASSIC REPTILE FOOTPRINTS 505

Brunswick fauna and its ecological relationships can be formu-
lated from the known footprints and bones.

Faunal content. In the preceding attempts to determine the
zoological positions of the trackmakers, failure to achieve a
correlation between footprint and reptile has been due less often
to inadequacy of the footprint material than to lack of com-
parable skeletal material. This lack is twofold in origin. First,
the Triassic reptiles as known are largely a spectral crew of
footless bodies and disembodied heads which can be made whole
only by persistent and lucky collecting. As our knowledge of
these forms increases, closer comparisons with the tracks will no
doubt become possible. Second, and more fundamental than the
fi-agmentary preservation of individual taxa, is the fragmentary
and selective preservation of the original fauna. At present we
know more types of Brunswick tetrapods from their footprints
than from skeletal material. Aside from a dinosaur leg excavated
by laborers but never collected (Colbert, 1946, p. 231), the only
genera known are the metoposaurid amphibian Eupelor, the
procolophonid cotylosaur Ilypsorjnathus, the phytosaur Clepsy-
saurus, and the aetosaurid pseudosuchian Stegomus.

Other Brunswick footprint assemblages — e.g. from Sanatoga
and Collegeville, Pennsylvania, and Tumble Falls and White-
hall, New Jersey — which would augment the fauna have been
omitted here because of the unreliability of old identifications.
Figured specimens of Anchisauripus [" Otouphepus"] poolei
(Bock) and RhyncJwsauroides ["Orthodactylus"] howelli
(Bock) from Schwenkville, Pennsylvania, which Wilhelm Bock
has kindly made available for study are specifically indeter-
minate but appear to record a eoelurosaur and a lacertoid rep-
tile of sphenodontid aspect. The Schwenkville " Grallator cnnea-
tus" (Bock, 1952, pi. 45, fig. 1) is surely not that genus but
instead has the phalangeal pattern of Anchisauripus; it evi-
dently represents another variety of eoelurosaur.

Ecology. The Brunswick fauna is predominantly terrestrial
but includes representatives of the aquatic biotope in the form
of metoposaurs and phytosaurs. These were certainly fish-eaters
and presumably fed on the coelacanths, semionotids, dictyopy-
gids and palaeoniscids of the Newark lakes and streams. The
presumed sphenodontid rhynchocephalians of Level B may (like

506 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Homoeosaurus) also have fed occasionally on fish, but the bulk
of their diet (like that of Sphenodon) probably consisted of
insects, other arthropods, grubs, and small tetrapods. Except
for the cotylosaur Hypsognathus, a presumed herbivore, the
other known members of the fauna were carnivorous. Lacertoids
and procolophonids furnished food for small, agile pseudo-
suchians of the Chirothcrium lulli type, and all three groups
formed the prey of the raptorial coelurosaurs. Whether the
larger Milford pseudosuchians were predators or carrion-feed-
ers, or both, cannot yet be determined.

Any assessment of the Brunswick ecology must allow for a
disproportionate percentage of aquatic animals in the skeletal
record ; the footprint horizons, on the other hand, record a more
representative proportion of terrestrial genera. More important,
the footprint faunules constitute biocoenoses in a strict sense,
while the skeletal remains occur as isolated finds or (at best)
as bonebed thanatocoenoses of animals which may or may not
have formed natural communities in life. Thus we can say with
confidence that the reptiles of Level B, for example, constituted
an animal community which occupied the Milford mudflat dur-
ing a period measured in days. Our sample is too small, un-
fortunately, to furnish significant data on the numerical com-
position of the faunules.

Survey of the Newark tetrapods. Undoubtedly the assemblage
described above is only a fraction of the actual tetrapod fauna
of Brunswick time. To afford a clearer perspective, the known
terrestrial vertebrates of the entire Newark pro^'ince are listed
with the Milford trackmakers in Table 4.

How incomplete this faunal picture remains may be deduced
from the mowj reptiles and amphibians, both genera and higher
groups, which are known from the Upper Triassic of Arizona,
Utah, New Mexico, Texas and Wyoming. Among these are the
metoposaurid amphibian Eupelor [^Bucttneria, etc.] (Colbert
and Imbrie, 1956), the large protorosaur Trilophosaurus, a vari-
ety of phytosaurs, the armor-plated pseudosuchians Desmatosu-
chus and Typothorax, the little ornithosuchid pseudosuchian Hes-
perosuchus, the coelurosaur Coelophysis, and the dicynodont Pla-
cerias. Many of these forms (or their relatives) must have in-
habited the Newark depositional area. Still other types of

TRIASSIC REPTILE FOOTPRINTS 507

tetrapods yet unknown, small forms in particular, completed the
fauna.

It would obviously be premature to attempt here a reconstruc-
tion of late Triassie tetrapod faunas in space and time on the
basis of skeletal and ichnological data now available. This is a
major project to which several researchers are individually con-
tributing, and in which the collaboration of invertebrate paleon-
tologists, paleobotanists, and geologists is essential. Nevertheless
the present study may serve to illustrate the contribution which
one field of investigation, ichnology, can make to the solution of
the general problem.

APPENDIX I: THE NATURE OF OTOUPHEPUS

A study of the type specimen of Otouphepus magnificus Cush-
man, lent through the courtesy of the Museum of Science, Bos-
ton, suggests that it is not the direct imprint of a foot but rather
an impression transmitted to a surface some millimeters below
the one on which the animal walked. This circumstance has
softened the outlines and produced the illusion of a compact
foot in which the digits were firmly united and surrounded
by a web. The dark-colored web outline described by Cushman
(1904, p. 155) proves to be a thin coat of gum which was readily
removed with soap and water. Analysis of the pattern of articu-
lar pads reveals a pedal structure which is well within the limits
of Anchisaiiripus. A previously undescribed claw-mark occupies
exactly the position of an Anchisauripus hallux imprint.

The only plesiotype cited by number (Lull, 1953, p. 177) is
morphologically indistinguishable from Anchisaurus tuberosus
— a fact which Lull has pointed out in recent correspondence.

In size and skeletal structure the unique specimen of 0.
minor Lull (1915, p. 190) is a typical Grallator; its peculiarities
are readily explained as artifacts of impression. (Latex molds
of the two preceding specimens were kindly furnished by the
Yale Peabody Museum.)

A third species, 0. poolei Bock (1952, p. 407) has been based
on an obscure and isolated footprint from the Brunswick forma-
tion near Schwenkville, Pennsylvania. With Wilhelm Bock's
gracious permission this specimen has been latex-molded and

508

BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY

MILFORD

TAXA

SKELETAL REMAINS

TRACKMAKERS
(as genera) (B)

AMPHIBIA

Stereospondyli:

Metoposauridae

Eupelor [Calamo'ps'] (S, G)
Dictyoceplialus (DR)

REPTILIA

Cotylosauria :

Procoloplionidae

Eypsognathus (B)

Rhynchocephalia (?)

2 fsphenodontids

Pseudosuehia:

2 large pseudosuchians

Ornithosuchidae

Stegomosuchus (P)

1 ?+ ornithosuchid

Aetosauridae

Stegomus (B, NH)

Phytosauria

Clepsysaurus (L, B. G, PD, NH)
Eutiodon (DR. ?G)

1 cf . Clepsysaurus

Coelurosauria :

3+ coelurosaurs

Ammosauridae

Ammosaurus (P)

Podokesauridae

Podokesaurus (P)

1 cf . Coelophysis

Prosauropoda:

Thecodontosauridae

Yaleosaurm (P)
Thecodontosaurus (L. P)

Cynodontia

Dromatherium (DR)
Microconodon (DR)

INCEETAE SEDIS

Gwyneddosaurus (L)

Lysorocephalus (L)

TRIASSIC REPTILE FOOTPRINTS 509

analyzed. An apparent structural formula of B, B, C, C allies
this species with the Anchisauripus tuherosus-exsertus-parallelus
species group. The species is probably distinct, but in the absence
of adequate material its small size is the only characteristic
which cannot be attributed to impression phenomena.

If Ofouphepus Cushman (March, 1904) and Anchisauripus
Lull (April, 1904) are synonymous the former takes priority.
To abandon a well-established name which has been applied to
several species comprising thousands of specimens, in favor of
a name which has been applied to four specimens whose chief
point of similarity is their obscurity, would be unreasonably
legalistic. I have therefore (Baird, 1956) petitioned that the
International Commission for Zoological Nomenclature exercise
its plenary powers to suppress Ofouphepus magniflcus as a
nomcn duhium ; this action automatically entails the suppression
of Lull 's family Otouphepodidae. At the same time Anchisauri-
pus sillimani and the family name Anchisauripodidae are to be
placed on the official lists of names in zoology as nomina con-
servanda.

APPENDIX II: BRONTOZOUM

The generic name Broniozoum, formerly applied to a number
of species of theropod footprints from the Newark series, was
erroneously reduced to synonymy in Hay's 1902 bibliography.
In his monograph of 1904 Lull followed Hay's usage, and since
that time the name has been in disuse. Broniozoum nevertheless
remains a valid prior generic name for the type species of Anchi-
sauripus Lull, and could legally be revived to replace that well-
known generic name. The facts are as follows :

Broniozoum Hitchcock, 1847, was proposed to include "five"
species of which only three were cited by name : the new species

TABLE 4. SURVEY OF THE NEWARK TETRAPODS

Formations: Deep River coal measures (DR) of North Caro-
lina; Stockton (S), Lockatong (L), Brunswick (B) of Pennsyl-
vania-New Jersey basin; Gettysburg shales (G) of York County,
Pennsylvania (Wanner, 1926) ; bed below Palisades diabase
(PD) of New Jersey; New Haven (NH), Meriden, Portland (P)
of Connecticut Valley.

510 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

parallelum,^^ sillimmiium (an invalid emendation of silUmani,
1843) and giganteuni. No type species was designated. No sub-
sequent author, to my knowledge, has designated a type species,
so all three remain available.

A case of sorts could be made for the selection of parallelum
through a broad construction of Article 30 A(a)(ii) of the
International Rules of Zoological Nomenclature, which provides
that the expression "n.g., n.sp.," can constitute a designation
of type species. In this case such a construction is highly dubi-
ous, particularly as it cannot be claimed to represent the intent
of the original author : for Hitchcock evidently had no intention
of designating a type species for any of his footprint taxa.
Furthermore, parallelum is based on two specimens — neither
designated as type — which are not conspecific. My analysis
assigns one of these (Hitchcock's fig. 2 b) to Anchisauripus silU-
mani and the other (fig. 2 a) to Grallator tenuis. To avoid
confusion I designate the specimen represented by fig. 2 b as the
type of parallelum, and reduce this species to subjective syn-
onymy with sillimani.

The second species, Brontozoum sillimani, is the type species
of AnchisaMripus Lull, 1904 (see Baird, 1956, paragraph 6).
Lull clearly erred in erecting a new genus with a type species
for which the name Brontozoum was available.

The third species originally assigned to Brontozoum, B. gigan-
teum, was first described by Hitchcock (1836) as a species of
Ornithichnites and was later transferred by him to Euhrontes
(1845) and to Brontozoum (1847). In 1902 Hay designated it
as the type of Euhrontes.

Since according to Article 30 B(g)(i) of the International
Rules the designation of a species as the type of one genus does
not in itself preclude the selection of that species as the type
of another genus, either sillimani or giganteum is eligible to be
the type species of Brontozoum. If sillimani be selected then
Anchisauripus (1904), an important name accepted for more
than fifty years, must be replaced by Brontozoum (1847), a
name generally rejected or ignored during the same period. If
giganteum be selected, however, then Brontozoum becomes a

12 This species B. parallelum is not to be confused with Anchisauripus [Gralla-
tor] parallelus (Hitchcock, 1865) although Hay (1902, p. 545) did so confuse it.

TBIASSIC REPTILE FOOTPRINTS 511

junior objective synonym of Eubrontes (1845) through the iden-
tity of their type species.

I therefore designate Ornithichnites giganteus Hitchcock, 1836,
as type species of the nominal genus Brontozoum, thereby syn-
onymizing that genus with Eubrontes. This suppression of
Brontozoum serves to maintain the familiar nomenclature of
Lull's classic monographs of 1915 and 1953 which have done so
much to bring nomenclatural stability into a confused field.

SUMMARY

Problem and method. Quarries in the Delaware valley near
Milford, New Jersey, have yielded three faunules of reptile
footprints at three horizons of the upper Brunswick formation
(Newark series, Upper Triassic). Each determinable species
was analyzed by comparison of outlines printed from latex molds
of all adequately preserved specimens. Body proportions and
locomotor habits of the traekmakers were deduced from track-
ways (where available) and the pedal skeletons were recon-
structed from the arrangement of articular swellings in the
footprints. Comparison with other Triassic footprints furnished
data on faunal distribution and stratigraphic correlation; com-
parison of the restored skeletons with those of Triassic reptiles
served to elucidate and enlarge the known Brunswick fauna.

Reddish-brown siltstone layer. The youngest faunule (dis-
cussed in a previous paper) records three reptilian genera as
footprint species : an indeterminate small dinosaur, another re-
sembling Coelurosaurichnus (sensu stricto) of the German Mid-
dle Keuper and the Gettysburg shale of Pennsylvania, and an
ornithosuchid pseudosuchian, Chirotherium lulli Bock, the last-
known member of its genus. This faunule and that of Level B
are the only American Chirotherium-dinossinr associations
known.

Level A (gray sandstone over shale). Grallator sulcatus, n.
sp., a small dinosaur with digits II-III united and IV inde-
pendent, is less digitigrade than its Connecticut Valley con-
geners. Among eoelurosaurs the massive German Halticosaurus
(as shown by pedal reconstruction) made footprints resembling
Eubrontes; little Procompsognathus compares more closely with

512 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Grallator, especially G. cursorius; and Coelophysis correlates
approximately with G. stdcatus.

Level B (gray sandstone over shale). Three dinosaur genera
occur: Anchisauripus milfordensis (Bock), a broad -based foot-
print without functional hallux, found also in the Gettysburg
shale of York County, Pennsylvania; A. parallelus (Hitchcock),
only 6/11 the size of conspecific New England footprints; and
an enigmatic small form of York County affinities. Taxonomie
placement follows a new type of key l)ased on relative posi-
tions of phalangeal and metatarso-phalangeal joints. Re-analysis
of various TTpper Triassic theropod tracks by this method
necessitates reassignment of several species and emphasizes the
similarities between Grallator and Anchisauripus (sensu stricto),
genera previously considered familially distinct but here united
as Grallatoridae and correlated with coelurosaurian dinosaurs.

Two large chirotheriids with hoof-like fifth pes digits represent
a lineage previously known only from the German Middle Keu-
per, here designated the brachychirotherian group. In Chiro-
therium parvuni (C. II. Hitchcock) (=C. copei Bock), narrow
claws on pes digits I-III were carried well above the distal pad
and directed obliquely laterad ; pes digit I was robust. In the
new species C. eyerniani straight, hea\y claws I-IV lay at sole
level and digit I was weak. Skeletal reconstructions of the large-
manus C. harthii (cosmopolitan. Lower Triassic) and the small-
manus C. lomasi (Keuper of England) contrast with those of
the Milford brachychirotherians. All three Milford chirotheriids
show parallel evolutionary tendencies — characteristic of late
Triassic species — toward functional tetradactyly through elim-
ination of lateral propping function in pes digit V. Inefficient
locomotion and increased dinosaurian competition probably con-
tributed to the extinction of Chirotherium. Pseudosuchians cor-
relative with the large Milford species are unknown.

Discovery of additional material permits redescription of
"Otozoiim" lineatus Bock as a new type of footprint, Apatopus
n. gen., correlated with a phytosaur such as Clepsysaurus — the
first phytosaur footprints known. Crocodilian in trackway pat-
tern except for the forward-turned manus, Apatopus differs
from all crocodilians in its long fourth and functional fifth pes
digits. Webs apparently connected the claw baises. Both adults

TRIASSIC REPTILE FOOTPRINTS 513

and juveniles are represented, one adult having a gleno-acetabu-
lar length of 52 cm.

Lacertoid footprints of the genus Rhynchosauroides [Akro-
pus'\ occur in the Brunswick, Gettysburg, and Lockatong forma-
tions of Pennsylvania-New Jersey, the Dockum of New Mexico,
and the Keuper of England and Italy. Rhynchosauroides hyper-
hates, n. sp., represents an agile reptile some 75 cm. long with
a gleno-acetabular length of 18 cm. Pes regularly overstepped
manus, and both were digitigrade with the metapodio-phalangeal
joints carried clear of the ground. A smaller Milford species
is inadequately known. Affinities appear to lie with the spheno-
dontid Rhynchocephalia or the Lacertilia.

Correlative age. The Milford faunules are akin to those of the
Pennsylvania-New Jersey basin and the German middle to late
Middle Keuper but not (as sometimes held) to the familiar foot-
print assemblages of the Portland and Meriden formations of
New England. Only one of the dinosaurian species also occurs
in the Connecticut Valley; while Chirotherium, Apatopus, and
Rhyncliosauroides are definitely absent from the abundant and
thoroughly studied Portland and Meriden faunas. No topo-
graphic or ecologic barriers would seem to have prevented inter-
regional migration. Presence of the phytosaur Clepsysaurus and
the pseudosuchian Stegomus in the lowest formation (New
Haven arkose) of the Connecticut Trias as well as in the Bruns-
wick formation, and the possible correlation of the three episodes
of igneous extrusion in New Jersey with the three lava flows of
Meriden time, suggest that the Brunswick antedates the Port-
land. The Milford footprint beds of the upper Brunswick ap-
pear to correlate best with the New Haven arkose.

Reptiles of the Brunswick formation. Known skeletal remains
include undetermined dinosaurs, the cotylosaur Hypsognathus,
the aetosaurid pseudosuchian Stegomus, and the phytosaur
Clepsysaurus. To this fauna the Milford footprints add three or
four types of coelurosaur, one small ornithosuchid and two large
undetermined pseudosuchians, a phytosaur, and two lacertoids.
From these combined faunal lists the ecological picture may at
least be glimpsed, though only a fraction of the Brunswick fauna
is yet known. The present study serves to illustrate the possible
contribution of ichnology to the solution of Triassic faunal and
stratigraphic problems.

514 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY

Appendix I. The nominal genus Otouphejms proves to be a
catch-all for obscurely preserved dinosaur footprints. Its sup-
pression as a nonien dubium is proposed in order to prevent
possible displacement of Anchisauripus as a junior synonym.

Appendix II. The long-disused name Brontozoum also threat-
ens the stability of the later name Anchisauripus. Brontozoum
is therefore cashiered by the designation of a type species which
is also the type species of the earlier genus Euhrontes, so that
the genera become objectively synonymous.

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TRIASSIC REPTILE FOOTPRINTS 517

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PLATES

PLATE 1

Fig. 1. Grallator sulcctus Baird, n. sp. Type, MCZ 215, a left pes imprint.
(As all specimens are natural casts, left and right appear reversed.) Fig. 2.
Chirotherium i)arvum (C. H. Hitchcock). Left manus-pes set, a composite
of MCZ 209 (pes) and MCZ 210 (manus).

b'

i'i-f

cm.

10

_!*;- «>

PLATE 1

PLATE 2

Chiro1]n liiim eiitrmani Baird, ii. sp. Three views of the type, MCZ 134,
a left pes ini])rint. Xote the fnnows along the lateral Ijorder of hoof-like
digit V.

r

<

TLATE

Apdtopii.s liiirdtiis (Boek). Fig. 1. Eight mnnus-pes set of the type indi-
vidual, MCZ •212. Pes digits IV and V failed to impress. Fig. 2. Well-
preserved left pes imprint, MCZ 213. Obliquely left of claw II is a manus
print of Ithyncliosaurnidrs sp. (cf. Figure 9 C).

PLATE i

Apatopus Uneatus (Boek). Fig. 1. Juvenile left iiianus-pes set, MCZ
214; note the beaded squaniation. Fig. 2. Juvenile left pes imprint,
AMNH 2259, rotated after implantation. Fig. 3. Small right pes imprint,
AMNH 2258.

lJ^f!l?fi.fliiiiliii¥r[Wb^iy

4::a.^^ -.. ■'■, ..^

PLATE 4