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UNIVERSITY OF KANSAS PUBLICATIONS
University of Kansas Science Bulletin - Vol. XXXIII - Part I

April 20, 1949
Lawrence, Kansas

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Science Bulletin

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No.

Contents of Volume XXXIII, Part I

Page

1. A Monograph of the Genus Taphrina A. J. Mix, 3

2 A Preliminary Account of the Herpetology of the State of

San Luis Potosi, Mexico : .Edward H. Taylor, 169

3. A Review of the Lizard Genus Barisia J. A. Tihen, 217

4 Costa Rican Frogs of the Genera Centrolene and

Centrolenella Edward H. Taylor, 257

5 Two New Teiid Lizards from Costa Rica

Edward H. Taylor, 271

6. New Salamanders from Costa Rica. . .Edward H. Taylor, 279

MUS. COM.P. ZOOL
LIBRARY

MAY 16 19^4^

HARMHO

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. I]

April 20, 1949

[No. 1

A Monograph of the Genus Taphri?ia

A. J. MIX

Abstract: The genus Taphrina, founded by Fries in 1832 on the species
'Taphrina populina, is the only recognized genug in the family Taphrinaceae,
order Taphrinales of the Ascomycetes.

All species of the genus are parasitic on higher plants or ferns, forming
mycelium (a) intercellularly ; (b) subcuticularly ; or (c) within the epidermal
wall ; forming asci in a subcuticular la5'er or in a wall locule ; overwintering in
the form of blastospores derived from ascospores by budding or in a few species
as perennial mycelium. Infection (so far as known) is by blastospores.

Asci arise from rounded ascogenous cells (chlamydospores), either by elonga-
tion of the ascogenous cell or by bursting out from the ascogenous-cell wall.
In many species a stalk cell (basal cell) is cut off from the ascus proper. Bud-
ding of the ascospores to form blastospores may occur within the ascus and
continues after spore expulsion.

Mycelium is dicaryotic. Fusion of nuclei occurs in the ascogenous cell;
meiosis in the young ascus. In one species (T. epiphylla) conjugation of
ascospores (or blastospores) occurs, resulting in dicaryotic hyphae. In other
species (so far as known) the dicaryotic condition is attained by division of
the single nucleus of the blastospore.

Species of Taphrina grow readily in artificial media if cultures are originated
from ascospores or blastospores; behaving in media as yeasts. Cells formed
in culture are blastospores, hyphae, ascogenous cells, and (rarely) asci.

Ninety-eight species of Taphrina are here redescribed and redefined (other
species being reduced to synonymy or excluded). Tliese are distributed by
hosts as follows: On Ferns, 24 species; on Populus and Salix, 4; on Betulaceae,
23; on Fagaceae, 5; on Ulmaceae, 2; on Rosaceae (except Prunus), 7; on
Pmnm, 17; on Rhus, 1; on Acer, 11; on Aesculus, 1; on Sebastiana, 1; on
Zingiberaceae, 2.

(3)

The University Science Bulletin

INTRODUCTION

IN earlier papers on the genus Taphrina an account of important
literature (Mix, 1936) and a list of presumably valid species
(Mix, 1936a) was presented. Since this species-list was compiled
from the literature and was not based on the study of actual speci-
mens, its value could be only temporary.

Following publication of these papers, morphological studies^
have been completed of all known species of Taphrina, and it is
now possible to undertake a thorough revision of the genus.

In certain cases the results of inoculation-experiments would be
helpful in reaching decisions on the identity of species, but for the
most part delimitation of species can be made on morphological
grounds. Interesting as biologic relationships will prove, their
elucidation is not likely to result in either combination or division
of morphologically evident species. Therefore, although many
species of Taphrina have been obtained in culture, and host rela-
tionships are being studied by means of inoculations, it seems un-
necessary to await the results of these studies before proceeding to
a revision of the genus.

The present paper gives an account of all known species of Taph-
rina. It is proposed to treat these species by host groups, discuss-
ing first species occurring on ferns and then those on Dicotyledons
and Monocotyledons, following the taxonomic arrangement of host
genera. For nomenclature of woody plants Rehder's Manual of
Cultivated Trees and Shrubs has been consulted. Other higher
plants have been named in agreement with Index Kewensis and
ferns in accordance with the usage of the Gray Herbarium.

In this revision an effort has been made to disturb existing situ-
ations as little as possible, especially as regards division of species
and erection of new species therefrom. In some cases description
of new species has seemed unavoidable, and, on the other hand, it
has sometimes seemed necessary to combine two or more existing

1. These studies were made, for the most part, during the period 1935-1940. They were
aided by mycologists the world over who sent valualile specimens as loans or gifts. Space
will not permit naming all of these collaborators but grateful acknowledgment to them is
hereby made.

Thanks are due also to the Chancellor and Board of Regents of the University of Kansas
for leave granted in 1939-1940, to Dr. Carl Hammarlund for laboratory facilities at Statens
Viixtskyddanstalt, Stockholm; to Dr. Th. Arwiddson for working privileges at the Botaniske
Riksmuseum, Stockholm; and to Dr. L. M. Massey for a temporary appointment in the
Department of Plant Pathology, Cornell University, during the winter, spring, and summer
of 1940.

The late Dr. D. H. Linder was so kind as to make available the facilities of the Farlow
Herbarium during the autumn of 1939.

Mix: Monograph of Genus Taphrina 5

species into one. Many species of Taphrina have in the past been
described as new without due regard to existing species. In some
cases occurrence on a new host was considered sufficient reason for
erecting a new species. In other cases morphological peculiarities
described by the author of a species have been found to disappear
when a number of specimens are examined. Finally, in a few in-
stances, species were hastily and inaccurately described.

In determining the validity of species the principle followed has
been to treat as identical those forms that are morphologically
similar and occur on related hosts. This is considered sound pro-
cedure even when biological distinction is known to exist between
different host-forms. The well known treatment accorded the rust
fungi by workers within that group may be cited in defense of
this point of view.

In critical cases type specimens have been examined. This has
not been thought necessary in the case of old and well established
species, much studied by earlier authors, such as Taphrina populina,
T. deformans, T. pruni. Whenever possible study of the type speci-
men has been supplemented by examination of a number of addi-
tional specimens, which has invariably resulted in widening the
limits of ascus-size given by the original authors. Often a similar
correction has resulted from the study of the type specimen itself.

The species descriptions, then, presented in this paper are in most
cases revisions of the descriptions given by the original authors,
and, being based on the study of as many specimens as possible,
may be accepted as accurate descriptions of the fungi concerned
insofar as they are known.

Morphological features considered in delimiting species are those
used by all previous investigators: habit of mycelium (whether in-
tercellular, subcuticular, or growing within the host-cell wall),
shape and size of asci, presence or absence of a stalk cell, shape
and size of stalk cells, and (rarely) size of spores.

CHARACTER OF THE GENUS AND RELATIONSHIPS

The order Taphrinales {Exoascales) of the Ascomycetes is com-
monly held to contain but one family, the Taphrinacae (Exoascacae)
Gaiimann (1926) includes a second family, the Protomycetaceae,
with two genera: Protomyces and Taphridium. All members of the
Protomycetaceae are parasitic, causing galls within the tissues of
higher plants.

Species of Protomyces form in the tissues of their hosts large,
round, thick-walled, overwintering chlamydospofes. Such a chlamy-

6 The University Science Bulletin

dospore germinates in the spring, its outer wall rupturing and the
inner thin-walled cell emerging as a sporangium (ascus or synas-
cus). In the peripheral cytoplasm of the sporangium numerous
nuclei divide by meiosis (Biiren, 1915), each producing a tetrad
of spore mother cells, which by further division form the spores.
Spores, after expulsion, conjugate in pairs.

The genus Taphridiiini differs from Protomyces in that the
chlamydospores are formed in a continuous layer beneath the host
epidermis. All known species of Taphridium were originally de-
scribed as species of Taphrina (Magnusiella) .

In the family Taphrinacae one valid genus (Taphrina)^ is known.
The type species is Taphrina populina Fries. Fries first (1815)
called this fungus Taphria populina aurea and subsequently (1825)
changed the genus name to Taphrina, stating that Taphria had
been used as an insect name. In Systema Mycologicum (1832),
Fries used the name Taphrina populina for this fungus.

All known species of Taphrina are parasitic, though susceptible
(so far as investigated) of artificial cultivation in the asexual stage
(Klebahn, 1923, Mix, 1924, Wieben, 1927) 3. Cultures may be ini-
tiated by ascospores ^ or by blastospores derived from them. Slow
growing, yeast-like, pale pink colonies are formed on various media
and cultures can apparently be propagated indefinitely. Cells
formed in culture are chiefly blastospores (yeast cells or bud-coni-
dia), though occasionally short hyphae may be observed, and thick
walled ''resting cells" are common. These "resting cells" are be-
lieved by the writer (Mix, 1924, 1935) to be ascogenous cells like
those formed beneath the cuticle of the host. Martin (1940), not
finding binucleate "resting cells" in her cultures, believed them to be
vegetative cells. In old cultures dicaryotic hyphae (like those in
host tissues) may occur, as well as ascogenous cells with fusing
nuclei, and even imperfectly formed asci. (Mix, 1935.) Roberts

2. Other genera have been recognized: Ascomyces, Exoascus, Magnusiella, but since the
papers by Johanson (1886) and Giesenhagen (1895, 1901) the common practice has been to
unite these all into one genus, for which the name Taphrina has priority. A full account of
the use of these different generic names by various authors has been given earlier (Mix, 1936).

3. Following the discovery by Klebahn (1923) and by the writer (1924) that species of
Taphrina could be grown in pure culture, Martin (1925) published notes on cultural behavior
of several species. Examination of her cultures, made shortly afterwards, showed that three
of them were wrongly named. One, called by Martin Taphrina communis has proved to be
a species of Torulopsis. Her culture from Quercus nigra, called Taphrina caerulescens, is a
species of Rhodotorula, and another isolate so named but from Q. rubra is a second and
different species of Rhodotorula. Apparently the rest of Martin's cultures were authentic.

4. The writer's procedure in obtaining pure cultures is to place ascus-bearing leaf-frag-
ments in the cover of an inverted petri dish of potato glucose (or other) agar. Ascospores
are shot upward onto the agar, the spores of each ascus coming to lie together in a sym-
metrical group. If the process is not allowed to continue too long the groups of spores are
spaced well apart and resultant colonies can be observed microscopically until it is convenient
to transfer them.

Mix: Monograph of Genus Taphrina 7

(1946) considers that ascogenous cells and asci may be formed in
culture, but was unable to find any binucleate cells.

In nature the ascospores bud readily, either within the ascus or
after spore expulsion. There is reason to believe (Fitzpatrick, 1934,
Mix, 1935) that propagation by budding in this "yeast stage" may
continue indefinitely, the fungi surviving in this fashion on various
plant surfaces and probably also in the soil. In other words species
of Taphrina are yeasts (in the broad sense) during their asexual
cycle. In fact if a species of Taphrina were brought into culture
without knowledge of its previous parasitic existence, it would be
called a species of Torulopsis.

In some cases, as in Taphrina epiphylla, infection apparently
occurs soon after ascospore expulsion, but in other cases (presum-
ably because host organs are susceptible only when young and ten-
der), a period of oversummering and overwintering is undergone
by the blastospores before infection. The occurrence of the "yeast
stage" seems to be important in accomplishing survival during this
prolonged period apart from the host tissues. For example, it is
evident (Fitzpatrick, 1934, Mix, 1935) that Taphrina deformans
can survive as a yeast for more than one year.

Infection by blastospores has been observed by Fitzpatrick
(1934) and Mix (1935) in Taphrina deformans. Successful in-
oculation experiments were performed by Sadebeck (1882) with
Taphrina bullata and T. tosqidnetii and later (1888, 1890) with
T. epiphylla. Fisch (1885) also reports successful inoculations
with T. tosquinetii and T. epiphylla. Klebahn (1923), inoculating
from pure culture, obtained infection with Taphrina tosquinetii on
Alnus glutinosa, but failed in attempts with T. epiphylla, T. sade-
becki, and T. betulina on their respective hosts. Wieben (1927)
also used pure cultures in her inoculation-experiments, securing
positive results with Taphrina tosquinetii, T. epiphylla, T. de-
formans; negative with T. populina {T. aurea), T. bullata, and T.
klebahni.

Species of Taphrina are of three types as regards mycelial habit:
intercellular forms {Taphrina deformans, etc.) developing abundant
mycelium between the interior cells of leaf, stem, or fruit, and sub-
sequently forming a subcuticular layer of ascogenous cells; sub-
cuticular forms {Taphrina epiphylla, etc.) whose mycelium and
ascogenous cells grow only beneath the cuticle; and wall-inhabit-
ing forms {Taphrina laurencia, etc.), living entirely within the
outer epidermal wall of the host. In Taphrina maculans (Butler,

S The University Science Bulletin

c

1911) and T. linearis (Mix, 1939) this "wall-habit" is developed
further. Flat bands of hyphae occur within radial walls between
adjacent cells of the epidermis and of the underlying tissues. None
of the intercellular or subcuticular forms is known to possess
haustoria but specialized haustoria have been reported (Butler,
1911) for T. maculans, and unspecialized haustoria for T. laurencia
(Giesenhagen, 1892), T. osmundae and T. higginsii (Mix, 1947).
Most wall-inhabiting forms produce numerous ascogenous cells
within a wall locule (Mix, 1939) but in the two species just named
a single multinucleate hyphal segment occurs in each wall locule
and this later gives rise to a single ascus (Mix, 1947).

Intercellular mycelium of species of Taphrina is for the most
part developed within the spaces between host cells, but in passage
of an infection hypha inward through the epidermis, and (later on)
of mycelium outward to form a subcuticular layer, growth occurs
perforce within radial walls between epidermal cells. In discuss-
ing Taphrina deformans, Martin (1940) states: ". . . vertical
hyphae are found between radial walls of epidermal cells and not
in these walls as described by Mix." This is a contradiction in
terms. With no radially-placed intercellular spaces between epi-
dermal cells of the peach only one location is possible for the ver-
tically-growing mycelium and this may be described either as ''in
the cell wall" or "between cell walls." The writer's expression
was chosen since it is not known whether the mycelium of Taphrina
deformans dissolves out the middle lamella or whether it may not
also attack the cellulose membranes.

Several species of Taphrina have been investigated cytologically.
There is general agreement (Dangeard, 1895, Ikeno, 1901, Juel,
1921, Martin, 1924, Eftimiu, 1927, Martin, 1940) that mycelium
with the host tissues is dicaryotic, each cell containing one or more
pairs of nuclei which divide conjugately. Nuclear fusion occurs
in ascogenous cells, as reported by Dangeard (1895) for T. de-
formans and by Juel (1921), Eftimiu (1927), and Martin (1924,
1940), for this and other species. The ascogenous cell is termed
a chlamydospore by Juel.

In germination of the chlamydospore (ascogenous cell) two types
of behavior have been observed. Juel (1921) found that in Taph-
rina epiphylla and in T. sadebeckii a pore forms in the outer wall
of the chlamydospore, and the ascus emerges as a thin walled en-
dospore through this widening pore.

Mix: Monograph of Genus Taphrina 9

In other species: Taphrina betulina, T. carnea, Juel found that
the chlamydospore itself elongates to form the ascus; its wall be-
comes thinner in the process but does not rupture.

Martin (1940) writing of Taphrina deformans says: "After the
nuclear fusion, the ascogenous cell elongates vertically and its thick
wall is stretched to become the thinner wall characteristic of the
ascus. The wall of the ascogenous cell does not break for the ascus
to emerge as described by Pierce (1900) and Juel (1921)." Juel
did not discuss Taphrina deformans in the paper referred to but
Pierce (1900) writes as follows: "As already said, the walls of the
ascogenous cells are heavy. The early steps in the development of
the asci from these cells (the development of a papilla-like elevation
on the upper surface of the cells) cause the rupture or the disso-
lution of the heavy wall where the elevation occurs. The phe-
nomenon is that of the germination of a heavy-walled spore, or
perhaps, more properly, the outgrowth or prolongation of an en-
dospore through the rupture of the epispore."

Whether the ascus of Taphrina deformans ever forms in this
manner seems to be difficult to determine. In all preparations
made by the writer the ascogenous cell elongates to an ascus, in
the fashion described by Martin (I.e.). If Pierce's observations
are correct they cannot hold true for the majority of asci.

On the other hand, this phenomenon of ascus emergence by rup-
ture of the chlamydospore wall (as described by Pierce for Taphrina
deformans, and by Juel for T. epiphylla) is readily seen in certain
other species. It has been observed (as noted below) in Taphrina
tosquinetii, T. sadebeckii, T. amentorum, T. epiphylla, T. occi-
dentalis, T. thomasii, T. acerina and T. acericola.

Meiosis occurs in the young ascus (Juel, 1921, Eftimiu, 1927,
Martin, 1924, 1940). Eftimiu (1927) reports the diploid number for
several species as four, while Martin (1940) reports it for Taphrina
deformans as eight, Juel and Eftimiu find meiosis occurring in
the first division of the fusion nucleus, Juel stating that in species
possessing stalk cells the protoplasm migrates into the upper portion
of the ascus before nuclear division, and that later an empty stalk
cell is cut off. Martin (1924, 1940) reports that in T. coryli and
T. deformans, the first division of the diploid ascus nucleus is
mitotic and that the stalk cell possesses for a brief period a diploid
nucleus and cytoplasm. These observations of Martin will, if con-
finned, be extremely important, since the occm-rence of a diploid

10 The University Science Bulletin

thallus is rare among fungi, having been previously reported by
Guilliermond and others for certain yeasts (Guilliermond, 1940),
and by Guilliermond (1928) for Spermopthora gossypii. A close
relation between the Taphrinales and the Saccharomycetales would
be indicated.

Ascospores are uninucleate and haploid, as are the blastospores
derived from them. The dicaryotic condition seems to .be attained
in most species (Eftimiu, 1927, and others) by nuclear division, a
dicaryon thus being a pair of sister nuclei.

In Taphrina epiphylla {T. klebahni is, as will appear later, a
synonym) Wieben (1927) reported conjugation between sexually
different ascospores (or blastospores) and development of a dicary-
otic hypha from the fusion-cell. (Presumably this is the infection
thread but the point has not been determined.) Wieben's obser-
vations were confirmed by the writer (1935).

Conjugation has not been observed as occurring regularly in any
other species, though it seems to occur rarely in Taphrina defor-
mans (Mix, 1935). With this fungus, however, infection has been
obtained by Fitzpatrick (1934) and the writer (1935) using cul-
tures derived from single ascospores.

Mix: Monograph of Genus Taphrina 11

TAXONOMIC ACCOUNT

I. SPECIES ON FERNS

Most species of Taphrina inhabiting ferns possess slender clavate
asci. Giesenhagen (1895), calling this the "Filicina-type" of ascus,
made it the basis for dividing the genus into subgenera. In other
respects, especially in mycelial habit (Mix, 1939) species on ferns
are remarkably diverse. That the series on ferns exhibits a variety
of development not equalled by the species on any other host group,
may be one indication of the anticjuity ^ of the genus.

1. Taphrina athyrii Siemaszko

Taphrina athyrii Siemaszko, Bull. Miisee du Caucase 12:20-28. 1919.

Causing small (5 mm. or less in diameter), yellow to brown, un-
thickened spots on leaves of Athyrium filix-femina Roth and Dry-
opteris spinulosa (Muell.) Kuntze. The spots are margined by small
veins, though they often occupy more than one vein-islet. At ma-
turity of the asci both surfaces of the spot are covered with a
whitish bloom except for a narrow unwhitened margin.

Mycelium subcuticular.

Asci amphigenous, broadly clavate to nearly oblong, rounded or
truncate at the apex, provided with a stalk cell. Ascospores round,
ovate, elliptic, or fusiform, frequently budding in the ascus. (Figure
1,A,B.)

Dimensions: of asci, 13-17[xX5.5-10[x of stalk cells, 4.5-8[ji,X
5-8[ji,; of spores, 2.5-6iJi, X 2-3.5[jl.

Distribution: On Athyrium filix-femina, Norway. On Dryopteris
spinulosa, Caucasus.

Material examined: A. filix-feynina. Norway: Sogn and Fjordane,
Hauglund, Brekke in Lavik, Aug. 12, 1927, T. Lillefosse (received
from Ivar J0rstad) .

D. spinulosa. Caucasus: Pschu, Sept. 17, 1917. (Part of Sie-
maszko's type collection. Two lots of this were studied, one ob-
tained from Dr. A. E. Jenkins, one from Siemaszko.)

This fungus was described by Siemaszko as occurring on Athyrium
filix-femina, but in a letter accompanying a portion of his type

5. The occurrence of fossilized Taphrina amentorum in interglacial deposits near Ejstrup,
Denmark, is reported by Lind (1913).

12

The University Science Bulletin

Fig. 1. Asci (X9(X)) of, A, Taphnna athyrii on Dryopteris spinulosa;
B, on Athyrium filix-femina; C, T. hiratsukae on Onoclea sensihilis; D,
on Pteretis nodulosa; E, on Pteretis strulhiopteris ; F, T. blechni; G, T.
cystopteridis ; H, I, T. calif ornica; J, T. cornu-cervi.

Mix: Monograph of Genus Taphrina 13

material he stated that he had been mistaken and that the host was
Dryopteris spinulosa. The species name is saved from being a mis-
nomer by the Norwegian material. Taphrina athyrii is very close
to T. hiratsukae in all respects, but may be distinguished by the
consistently broader ascus and by the nearly isodiametric stalk cell.

2. Taphrina hiratsukae Nishida

Taphrina hiratsukae Nishida, Miyabe Festschrift, Tokyo, 1911.

T. struthiopteridis Nishida, 1. c.

T. struthiopteridis Siemaszko, Bull. Musee du Caucase, 12:20-28. 1919.

T. siemaszkoi (Siem.) Mix, Univ. Kansas Sci. Bull. 24:150-176. 1936.

Causing small (up to 5 mm. in diameter) , yellow (becoming brown
with age), unthickened spots on leaves of Onoclea sensibilis L.,
Pteretis nodulosa (Michx.) Nieuwl., Pteretis struthiopteris (L).
Todaro, and Thelypteris thelypteris (L.) Nieuwl. The spots may
be margined by the small veins of the leaf, though occupying more
than one vein-islet. At maturity of the asci the lower surface of
the spot becomes covered with a whitish bloom except for the nar-
row unwhitened margin.

Mycelium subcuticular.

Asci hypophyllous, clavate, rounded or truncate at the apex,
provided with a stalk cell. Ascospores ovate or elliptic, frequently
budding in the ascus. (Fig. 1, C, D, E.)

• Dimensions: Of asci, 13-30[x X 4-18[x; of stalk cells, 4-lOpL X
3-5[jl; of spores, 2-6|jl X 2-4tj..

Distribution: On Onoclea sensibilis. New York, Ontario, Penn-
sylvania, Japan. On Pteretis nodulosa, Manitoba, Wisconsin. On
Pteretis struthiopteris, Caucasus. On Thelypteris thelypteris,
Japan.

Material examined: Onoclea sensibilis. New York: Hudson Falls,
Aug. 7, 1919, J. Dearness. Pennsylvania: Houserville, Aug. 10,
1921, C. R. Orton and W. A. McCubbin. Ontario: swamp east of
Wilcox Lake, Aug. 4, 1930, H. S. Jackson. Japan: Pref. Iwate,
Morioka, July 22, 1934, K. Togashi.

Pteretis nodulosa. Wisconsin: Weyerhaeuser, J. J. Davis.

Pteretis struthiopteris. Caucasus: Pschu, Sept. 13, 1917, W.
Siemaszko (duplicate of Siemaszko's type material of T. struthiop-
teridis).

Thelypteris thelypteris. Japan: Pref. Iwate, Mt. Iwate, Aug. 17,
1903, G. Yamada.

In the species list previously published (Mix, 1936a), Taphrina
struthiopteridis Siemaszko was, on the basis of Siemaszko's descrip-

14 The University Science Bulletin

tion, held to be distinct from T. struthiopteridis Nishida, and the
fungus was therefore renamed Taphrina siemaszkoi. Siemaszko de-
scribed this fungus as having asci 28-40pL X 6-7[x and did not men-
tion a stalk cell, while Nishida's fungus was described as having
asci 16-28pL X 4-6[j(, with stalk cells 4-7[x X 3-4[jl. In a letter accom-
panying his type material Siemaszko stated that his measurements
had been wrong, giving the following as correct: Asci 17.5-32[jl X
6-7.5[jL, again not mentioning the stalk cell. He said further that
his fungus was probably identical with T. struthiopteridis Nishida.
Study of the material obtained from Siemaszko, and of duplicate
material in the Farlow Herbarium showed asci 17-28[jl X 5-7ij., and
stalk cells 4-7ii. X 3-6pi..

It is not possible to distinguish from each other the fungi under
discussion either on the basis of host lesions or of size and shape
of asci and stalk cells. They are therefore held to be the same
species.

Since Nishida described his two fungi in the same paper it be-
comes a question which name should receive priority. The name
Taphrina hiratsukae Nishida is chosen since it would precede T.
struthiopteridis in an alphabetical list of species. The close simi-
larity of Taphrina hiratsukae to T. athyrii has been mentioned
above. It is not impossible that these two species are identical.

3. Taphrina blechni (Bresadola) ex Mix.

Taphmia blechni Bresadola ex Mix, Trans. Kansas Acad. Sci. 50:77-83.
1947.

Causing small, round or ellipsoid, unthickened spots on leaves
of Blechnum sp.

Mycelium subcuticular.

Asci amphigenous, clavate, truncate at the apex, often with a
curved foot, lacking a stalk cell. Ascospores ellipsoidal. Numerous
elongate spores (apparently blastospores) may be present in the
ascus. (Fig. 1, r.)

Dimensions: Of asci, 23-60[jl X 4-7ijl; of ascospores, 2-5[J(, X 2-4^/.;
of blastospores, 5-6. 5ix X 0.5-1.5ij..

Distribution: Brazil.

Material examined: Brazil: Rio de Janeiro, Serra Geral, Oct.
1891, E. Ule (E. Ule, Herb. Brasil. 1786). Bot. Mus. Berlin, Bot.
Mus. Stockholm.

Bresadola named (in the above mentioned Exsiccati) this fungus
but did not publish a description. This has been supplied by the
writer (Mix, 1947a).

Mix: Monograph of Genus Taphrina 15

4. Taphrina cystopteridis Mix
Taphrina cystopteridis Mix, Mycologia 30:563-569. 1938.

Causing small (0.5 to 2.0 mm. in diameter) galls on leaves of
Cystopteris fragilis (L.) Bernh.

Mycelium intercellular.

Asci epiphyllous or amphigenous, clavate, rounded or truncate at
apex, provided with a stalk cell. Ascospores eight, round, ovate or
elliptic.

Dimensions: Of asci, 20-30[jl X 4-7. 5^;,, of stalk cells, 6-19[x X
4-6[x, of spores, 3-6 X 2-3.5[jl. (Fig. 1, G.)

Distribution: Indiana, Kansas, Wisconsin.

Material examined: Indiana: Daviess County, Glendale, June
4, 1923, C. C. Deam; Greencastle, May 23, 1922, id. Kansas:
Neodesha, June, 1936, W. H. Horr (type material) ; same locality,
June, 1937, and June, 1938, A. J. M. Wisconsin: Brodhead,
Sept. 16, 1926, J. J. Davis.

The earliest collection of this fungus is the one made at Green-
castle, Indiana, by Deam, and the earliest mention is by Davis
(1929) who identified the fungus collected at Brodhead, Wis., as
Taphrina filicina Rostr.

5. Taphrina californica Mix
Taphrina californica Mix, Mycologia 30:563-569. 1938.

Causing small to large (several centimeters in extent when
swollen) orange-brown galls on leaflets, midribs, and petioles of
Dryopteris arguta (Kaulf.) Wats. The galls are fleshy and gela-
tinous, becoming small, wrinkled, and hard when dry; much swollen
when wet. The gall originates from the epidermis and the mycelium
develops entirely within the outer walls of the surface cells ("epi-
dermal cells") of the gall. Ascogenous cells develop in a much
enlarged wall locule, and the mature asci burst forth and protrude
from this locule.

Asci clavate, truncate or rounded at the apex, provided with stalk
cells; ascospores usually eight, ovate to elliptic. (Fig. 1, H, I.)

Dimensions: Of asci, 23-46[jl X 7-8[jl; of stalk cells, 17-30[ji, X
5-7pt. ; of spores, 4:-5]x X 2-3[jl.

Distribution: California, Mexico, Oregon.

Material examined: California: Lake Phenix, Marin County,
Aug. 21, 1930, Victor Duran; same locality, Sept. 4, 1930, H. E.
Parks (California Fungi 343) ; Lee Bonar, Nov. 28, 1937 (type) ;
Carmel, Big Sur River, June 23, 1938, C. E. Scott; Palo Alto, San

16 The University Science Bulletin

Francisquito Creek, Sept. 25, 1938, R. H. Thompson; San Mateo
County, Los Altos, May 30, 1939, R. H. Thompson.

The collection by Parks was widely distributed (as California
Fungi 343) under the name Taphrina filicina Rostr. Many sub-
divisions of this collection in various herbaria have been exam-
ined, but in all of them the fungus is immature, showing ascogenous
cells but no asci. The galls are characteristic.

Taphrina californica is apparently common in the coastal region
from lower California to central Oregon. Dr. R. H. Thompson
kindly searched the collection of ferns in the Stanford University
Herbarium, looking for those with unmistakable galls of T. cali-
fornica. Ferns bearing galls were found to have been collected
from the following localities: California: Amador County, Cedar
Creek; Marin County, Petaluma; Monterey County, above P. I.
Co. Dam; Rand County, City Creek; San Diego County, East Illi-
nois River, four miles from Takilma, seven miles above Lakeside;
San Mateo County, Santa Cruz Mountains, King's Mountain;
Santa Clara County, Los Gatos; Tulare County, Sequoia National
Park. Oregon: Multnomah County, Elk Rock. Mexico: Santo
Tomas (Baja California).

6. Taphrina cornu-cervi Giesenhagen
Taphrina cornu-cervi Giesenhagen, Flora 76:130-156. 1892.

Causing long, branched, antler-ljke outgrowths on leaves of
Polystichum aristatum (Forst.) Presl.

Mycelium intercellular.

Asci covering surface of outgrowth, clavate, rounded at apex,
provided with a stalk cell; ascospores eight, round, ovate or elliptic,
often budding in the ascus.

Dimensions: Of asci, 18-24ijl X 6-10[;,; of stalk cells, 5-8[x X 5-7[J.;
of spores, 3.5-4.5[x X 2-3ix. (Giesenhagen reported asci 24[a X 5-6[x,
stalk cells 4-6ix X 2-4[jl, spores not present.) (Fig. 1, J.)

Distribution: Ceylon, Fiji Islands, India, Japan, Nepal, New
Caledonia; Tahiti.

Material examined: Fiji: Nodarivatu, Viti Levu, May-July,
1927, H. E. Parks (Plants of V. L. F. Coll. by Bernice P. Bishop
Museum and Univ. Calif.) India: Nilgiri, no date, ex Herb.
Sydow (in Bot. Mus. Stockholm) ; Coonoor, Sim's Park, Nilgiri
Hills, no date (received 1939), M. 0. P. Iyengar. Japan: Hame-
mura, Prov. Tosa, 1908 (ex Herb. Morioka Imp. Coll. Agric. and
For.).

Mix: Monograph of Genus Taphrina 17

In addition to the preceding the following three specimens have
been examined superficially. They bear no dates. They are in
the Patouillard Herbarium, now owned by the Farlow Herbarium.
One is labelled: "Herbarium Stendel, PL Indiae Or. (M. Nilgiri)
Ed. R. F. Hohenacher, 901. Roestelia tubaeforniis Rabenh, n. sp.,
In Aspidio cervifolio Kze. In montibus Nilgiri." The second:
"Taphrina tubiforme Lagerh., Nouvelle Caledonie." The third:
"Taphrina tubiforme Lagerh., Tahiti. Herb. Rancher." In these
specimens the outgrowths are characteristic.

Giesenhagen (1892) described the fungus from material collected
in Nepal and Ceylon. Sadebeck (1895) reports specimens from
Queensland, Fiji, and Samoan Islands.

7. Taphrina vestergrenii Giesenhagen

Taphrina vestergrenii Giesenhagen, Bot. Zeit. 59:115-142. 1901.
Exoascus vestergrenii (Gies.) Saccardo and Sydow, S'ylloge Fungorum
16:1152.

Causing small (up to 5 mm. diameter), fleshy, brownish (prob-
ably lighter colored in fresh material) galls on leaves of Dryopteris
filix-mas Schott, and perhaps also on D. lacera 0. Kuntze.

Mycelium subcuticular.

Asci hypophyllous or amphigenous, clavate, rounded or truncate
at the apex, stalk cell pointed or tmncate below.

Dimensions: Of asci, 23-50pL X 6-10[jl; of stalk cells, 10-23[ji, X
4-lOpi,; of spores, 3.5-6ia X 2.5-4[jl. Ascospores eight, round, ovate
or elliptic, often budding in the ascus. The asci are closely packed
and in some specimens both asci and stalk cells show an irregular
contour. (Fig. 2, A.)

Distribution: Northern and Central Europe (Japan and China?).

Material examined: Denmark: Skjelskor, June 23, 1907, J. Lind.
Germany: Allgau, Einodsbach, Feldhom, July, 1909, W. Krieger;
Alsace, Hohneck, Fischboedle, July 14, 1910, H. Sydow (Sydow
Myc. Germ. 978) ; Baden, Feldhorn, Aug., 1903, G. Lagerheim.
Russia: Abro Island, near Osel, July 1, 1899, T. Vestergren (Rehm.
Ascomyceten, 1412, dupl. of type). Sweden: Uppland, Kuggvik,
Lilla Moja, July 4, 1923, T. Vestergren.

The asci were described by Giesenhagen as 25pL X 6[x, and dimen-
sions of the stalk cell were not given. Jankowska (1928) gave di-
mensions of asci as 24-35iJ. X 5-7[x, of stalk cells as 12-21iJi. X 4-6^1.,
and stated that stalk cells may be pointed. Spore-dimensions as
given by Giesenhagen and Jankowska are 7[jlX2.5-3[jl.

2—3659

18

The University Science Bulletin

Fig. 2. Asci (X 900) of A, Taphrina vestergrenii; B, T. gracilis; C,
T. jusca; D, T. jiiaca (Bubak's "T. morijormls'), ascogenous cells and
one ascus; E, T. filicina.

Besides these specimens of the undoubted T. vestergrenii three
specimens from the Herbarium of the Morioka Imperial College
of Agriculture and Forestry have been studied, all labelled "Taph-
rina vestergrenii Giesenhagen on Dryopteris lacera 0. Kuntze":
Japan: Prov. Ise, Mt. Komono, Aug. 1903, K. Nakanshiki; Prov.

Mix: Monograph of Genus Taphrina 19

Kochi, Mt. Yanaze, Oct. 1904, T. Yoshinaga; Prov. Tosa, Higi-
shikawamura, Aug. 1905, T. Yoshinaga. The galls in these speci-
mens are overmature, and no asci can be found. The galls show
greatly distorted vascular bundles, a feature not observed in the
European specimens.

A specimen of a fungus on an unidentified species of Dryopteris
collected at Mt. Omei, Szechwan, China, Aug. 3, 1928, by W. P.
Fang may be Tafhrina vestergrenii. The fungus is immature, show-
ing only subcuticular mycelium, but the gall resembles that on Dry-
opteris lacera. The specimen in question was kindly donated by
Dr. D. H. Linder, and a duplicate remains in the Farlow Her-
barium.

8. Taphrina gracilis Mix
Taphrina gracilis Mix, Mycologia 30:5:563-579. 1938.

Causing small (up to 5 mm. diam.) thickened areas, resembling
small lesions of peach leaf curl, on leaves of Dryopteris marginalis,
(L.) A. Gray.

Mycelium intercellular.

Asci epiphyllous or amphigenous, clavate, rounded at the apex,
provided with a stalk cell; ascospores eight, fusiform.

Dimensions: Of asci, 26-36[x X 4-6pt,; of stalk cells, 13-23[x X
3-4[j.; of spores, 5-8tJ. X 2-2.5^;.. (Fig. 2, B.)

Distribution: Labrador Lake, New York.

Material examined: Cornell Univ. Dept. Plant Path., New York
Bot. Card., Brooklyn Bot. Card., Fungi collected near Ithaca, N.
Y., June 2-7, 1919, No. 1256, Labrador Lake, Coll. E. W. Olive,
A. H. W. Povah, L. R. Hesler, F. J. Seaver, H. H. Whetzel, H. M.
Fitzpatrick, et al. In New York Bot. Gard. Herb. (Type). Dupli-
cate material at the other institutions named does not show the
fungus in mature condition.

Because this fungus is known only from a single meager collec-
tion the writer (1. c.) had some hesitation in describing it as a dis-
tinct species. The temptation was to consider it as a host-variant
of Taphrina fusca (known to occur in the same locality) or of T.
vestergrenii (not known in North America). However, the char-
acteristic slender asci and the fusiform spores differentiate it from
both these species. Moreover the gall is different from those formed
by T. fusca and T. vestergrenii (q. v.), being characterized by
moderate hypertrophy of all the cells of the lamina.

20 The University Science Bulletin

9. Taphrina fusca Giesenhagen

Taphrina jusca Giesenhagen, Flora 86:100-109. 1899.

Exoascus fuscus (Gies.) Saccardo and Sydow, Sylloge Fungorum 16:803.

Taphrina morijormis Bubak, Bull. Herb. Boissier, Ser. 2, 6:393-488. 1906.

Causing small (up to 5 mm. in diam.) fleshy (convoluted when
dry), yellowish galls on leaves of Dryopteris rigida (Hoffm.) Un-
derw., D. rigida var. australis Christ., D. spimdosa (Muell.) Kuntze,
D. spinulosa var. americana (Fisch.) Fern., and var. intermedia
Underw. According to Giesenhagen (1. c.) the gall, though exhib-
iting considerable hypertrophy, is epidermal in origin, and this
seems to be the case in the various specimens examined. Fresh
material, with galls of various ages, would be necessary in order
to determine this point with exactness.

Mycelium subcuticular.

Asci epiphyllous, or sometimes amphigenous, clavate, rounded or
truncate at the apex, provided with a stalk cell which may equal
or exceed in length the ascus proper. Ascospores globose to elliptic,
sometimes budding in the ascus. (Fig. 2, C, D.)

Dimensions: Of asci, 19-27[ji. X 4-8[jl; of stalk cells, 15-34[jl X
4-7[Ji; of spores, 3.5-liJ. X 2-3.5ijl.

Distribution: On Dryopteris rigida, Albania. On D. rigida var.
australis, Sicily. On D. spimdosa et var., eastern North America.

Material examined: Dryopteris rigida. Albania: Njegusi prope
Cetinje, July, 1903, F. Bubak (Bubak's type specimen of Taphrina
morijormis, obtained on loan from Brooklyn Bot. Gard.). D. rigida
var. australis. Sicily: Near Palermo, Nov. 1898, K. Giesenhagen
(Giesenhagen's type specimen of T. jusca, obtained on loan from
Univ. of Munich Herbarium). D. spinulosa et var. New Bruns-
wick: Campobello, July, 1902, W. G. Farlow (Rel. Farl. 149).
New Hampshire: Mt. Lafayette, July 20, 1935, D. H. Linder; Mt.
Washington, Tuckerman Ravine Trail, July 6, 1931, K. S. Chester
and J. H. Faull (Herb. J. H. F. 9934). Vermont: Mt. Mansfield,
July 15, 1932, J. H. Faull and K. S. Chester (Herb. J. H. F. 10734).
West Virginia: Bayard July, 1891, W. C. Sturgis.

Bubak's description of Taphrina morijormis gives dimensions of
asci as 30-45iJi. X 4-6ix, stalk cells not mentioned, spores not seen.
The asci are further described as narrow at the apex. Examination
of his type material makes it appear that he saw and described no
asci but only ascogenous cells. These are long and narrow, 30-46ijl X
5[x, and form a close layer beneath the epidermis (Fig. 2, D). Dili-
gent search of the material revealed a single mature ascus with a

Mix: Monograph of Genus Taphrina 21

stalk cell and spores. D;imensions of this ascus were 33 X 10[J., of
the stalk cell, 17 X 5pt., and of the spores, 4.5 X 5[ji.. The gall itself
is somewhat larger than in other collections of Taphrina fusca.
Judging from its dried state it may reach one centimeter in diam-
eter when fresh, and is much convoluted. That this convolution is
apparent in the fresh state may be inferred from Bubak's descrip-
tion and from his choice of a specific name. There is no doubt that
Bubak's and Giesenhagen's fungi, occurring on the same host spe-
cies, are identical.

The writer's reasons for assigning the American fungus on Dry-
opteris spinulosa to Taphrina fusca have been given earlier (Mix,
1938).

The yellowish or brownish coloration of the galls is found in the
walls of the outer cells ("epidermal cells"), all fungous structures
being hyaline.

10. Taphrina filicina Rostrup ex Johanson

Taphrina filicina Rostrup ex Johanson, Bih. Kongl. Vetensk. Akad. Handl.
13:3-28. 1887.

Ascomyces filicinus Rostrup, Vestergren, Micr. Rar. Sel. No. 813.
Exoascus filicinus (Rostr.) Saccardo, Sylloge Fungorum 8:819.

Causing thickened areas of the leaf-curl type or small fleshy galls
on leaves of Dryopteris spinulosa (Muell.) Kuntze, D. spinulosa
var. americana (Fisch.) Fern., and var. intermedia Underw. As
pointed out by Giesenhagen (1899) the gall is not epidermal in or-
igin, but all leaf tissues are involved.

Mycelium intercellular.

Asci amphigenous, clavate, rounded at the apex, attenuate at the
base, lacking a stalk cell. Ascospores ovate to elliptic. (Fig. 2, E.)

Dimensions : Of asci, 18-46[jl X 6-10[x; of spores, 3.5-6.5[x X 2-3. 5[j..

Distribution: northern Europe and eastern North America.

Material examined: Germany: Tirol, Sondestal, Geschnitztal,
Aug., 1939, F. Wettstein. Russia: Prov. Novgorod, Beresaida,
July, 1890, W. Tranzschel (Jaczewski, Komarov, Tranzschel. Fungi
Rossiae Exsiccati, 27). Sweden: Dalarna, Avesta, Elfnashage,
July, 1879, Conrad Indebetou (duplicate material from the type-
collection, Bot. Mus. Stockholm.) Poland: Krakow, no date, Raci-
borski (Ex Herb. A. Wroblewski). New York: Ithaca, Enfield
Gorge, July 14, 1917, J. H. Faull (Herb. J. H. F., 1804) ; ibid, July
14, 1917, L. B. Walker; Coy Glen, July 3, 1927, L. B. Walker; Mc-
Lean, McLean Preserve, July 25, 1937, W. W. Ray (Herb. W. W. R.
371). Pennsylvania: Mt. Jewett, McKean Forest, July 7, 1938,
A. E. Edgecombe and L. 0. Overholts (Ex. Herb. L. 0. 0.).

22 The University Science Bulletin

The wide range of variation in ascus size between European and
American specimens of this fungus has ah'eady been pointed out
(Mix, 1938). Many specimens of other species of Taphrina in
American herbaria are wrongly labelled T. filicina, American col-
lectors having referred fern-inhabiting species of Taphrina to this
species indiscriminately. Such specimens have been properly al-
located in the present account without, however, mention in every
case of the mistaken identification.

Jaczewski (1926) mentions Russian specimens of Taphrina filicina
on "Nephrodiiwi, Phegopteris, and Athyrium filix-jemina." It has
not been possible to obtain these specimens for study.

11. Taphrina thaxteri Mix

Taphrina thaxtcri Mix, Mycologia 31 :4: 445-454. 1939.

Causing pale yellowish unthickened spots, one centimeter or
more in extent, on leaves of Dryopteris poiteana (Bory) Urban.

Mycelium growing within the epidermal wall.

Asci hypophyllous, protruding from a wall locule, clavate, rounded
at the apex, provided with a stalk cell; ascospores eight, often
fusiform. (Fig. 3, A, B, C.)

Dimensions: Of asci, 20-27[j. X 5-7[jl; of stalk cells, 7-15[jl X
5-7pi.; of spores, 3.5-5[i, X 2-3[x.

Distribution: Trinidad.

Material examined: Trinidad: Arima: Verdant Vale, no date,
R. Thaxter (Herb. Roland Thaxter, ''Bequest 1932").

12. Taphrina fasciculata (Lagerh. and Sadeb.) Giesenhagen

Taphrina jasciculata (Lagerh. and Sadeb.) Giesenhagen, Flora 81:267-361.

1895.

Magnusiella fasciculata Lagerheim and Sadebeck, Ber. Deutsch. Bot. Ges.

13:265-280. 1895.

Causing small unthickened spots with a grayish or whitish ascus-
bearing center, and a narrow pale yellowish margin (thus resembling
spots caused by T. lutescens) on leaves of Dryopteris sp. [D. filix-
masl).

Mycelium intercellular.

Asci short-clavate and provided with a short stalk cell, or long-
clavate with a long stalk cell, or arising from a two-celled or three-
celled hypha. The longer asci with longer stalk cells are frequently
empty, having discharged their spores. The basal cell of a two-
celled or three-celled stalk often has a bent foot. Ascospores eight,
ovate to elliptic. (Fig. 3, D.)

Dimensions: Of asci, 20-53[x X 8-lOt^; of stalk cells, 10-66ia X

Mix: Monograph of Genus Taphrina

23

Fig. 3. A, B, C, Taphrina thaxteri; A, Ascogenous cells in wall locule;
B, Asci emerging from locule; C, Asci. D, Asci of T. jasciculata; E,
of T. ecuadorensis ; F, of T. osmundae. All X 900.

24 The University Science Bulletin

7-8[x; of second or third cells of septate stalk, ca. 40[x X 7[x; of
spores, 5-7[jL X 2.4-4[jl.

Distribution: Ecuador.

Material examined: Ecuador: Quito, Rio Machangara, no date,
Lagerheim, det. Sadebeck (Herb. Reg. Monac). Type specimen,
Munich Museum.

The inner packet containing this material bears the following,
presumably by Sadebeck: "Magnusiella Nephrodii Lagerh. in
herb., bildet kleine weissliche oder graiiliche nicht blasige fiecken auf
Nephrodium {filix-mas). Trotz vielem suchen nur an einen standort
und nur an einem wedel beobachtet. Der standort war sehr feucht
und schattig. Rio Machangara bci Quito, Leg. G. v. Lagerheim.
Am nachsten mit M. lutescens verwandt, hat jedoch lang keulige
asken mit eiformigen sporen (keine conidien), ahnelt daher M. po-
tentillae." Li another handwriting is added: "Magnusiella fasci-
cidata V. Lagerheim and Sadebeck."

Sadebeck gives dimensions of the asci as 50-70[x X 9-12[x, and
states that a stalk cell is present which is narrower than the ascus.
These dimensions are hard to understand unless Sadebeck included
stalk cell and ascus in his measuring.

It is impossible to determine the host species from the material
available.

13. Taphrina ecuadorensis Sydow
Taphrina ecuadorensis Sydow, Ann. Myc. 37:275-438. 1939.

Causing small (1.5 to 3 mm. diam.) yellowish to yellow brown
(with whitish central area bearing asci and narrow margin) un-
thickened or only slightly thickened spots on leaves of Dryopteris
cheilanthoides (Kze.) C. Chr.

Mycelium intercellular, abundant (stated to be subcuticular by
Sydow) .

Asci cylindric, clavate, rounded at apex, borne on the ends of sep-
tate hyphae, which often arise from a rather dense weft of mycelium
(in this habit resembling Taphrina potentillae) . The asci fonn
dense tufts, pushing up through the epidermis and rupturing it. As-
cospores eight, elliptic to fusiform. (Fig. 3, E.)

Dimensions: Of asci, 23-40[ji. X 6-10[x; of mycelial cell next to
ascus (corresponding to stalk cell), 23-56ix X 4-7[ji,; of spores, 4-6. 5[x
X 2-2.5[JL.

Distribution: Ecuador.

Material examined: Ecuador: Tungurahua: Hacienda San An-

Mix: Monograph of Genus Taphrina 25

Ionic pr. Bancs, Dec. 31, 1937, H. Sydcw (Fungi Aequatcriensis
651). Part cf type, obtained from Sydow.

Sydow wrote that he found the fungus "somewhat difficult to
study," which is indeed the case. He does not report the intercellu-
lar mycelium nor the rupturing of the host epidermis by the compact
tufts of asci, though he describes the asci as crowded ("dense con-
ferti"). Neither does he mention that the asci arise from the ends
of septate hyphae, but describes a stalk cell ("in matricem non
penetrante"), 20-35ijl X 4-6[i,.

This species is probably synonymous with the preceding but suf-
ficient material is not at hand to demonstrate it.

14. Taphrina osmundae Nishida
Taphrina osmundae Nishida, Miyabe Festschrift, Tokyo, 1911:157-212.

Causing small to large, yellowish to light brown, unthickened
spots (sometimes involving the whole leaflet), on leaves of Osinunda
regalis (L.) var. japonica Willd. A sparse white powdery layer of
asci covers the lower surface of the spot and less commonly smaller
areas occur on the upper surface.

Mycelium developing within outer walls of epidermal cells, form-
ing wall locules, each of which contains a single ascogenous cell.

Asci hypophyllous, less commonly also epiphyllous, scattered, one
ascus formed in each wall locule, emerging as a small papilla which
gradually enlarges to an oblong ascus, a small, triangular stalk cell
remaining inserted in the locule. Ascospores not seen, blastospores
numerous, often forming in the ascogenous cell (young ascus) before
ascus emergence; in mature asci the blastospores occur in a peri-
pheral layer close to the wall. (Fig. 3, F.)

Dimensions: Of asci, 26-63[jl X 17-27[jl; of stalk cells, 6-17[ji, X
8-17[jl; of spores (blastospores), 3-4[x X 2-3.5[jl.

Distribution: Japan.

Material examined: Japan: Pref. Tottori, Taisenji, July 1, 1924,
K. Togashi (Herb. Morioka Imp. Coll. Agric. and For. 193) ; Pref.
Iwate, Mt. Iwate, June 13, 1934, K. Togashi (Herb. Morioka Imp.
Coll. Agric. and For.) ; Kyusu, Mt. Kirisama, May 29, 1938, I. Hino
(Herb. Lab. Path. Veg. Miyazaki Koto Norni-Gukko).

In early stages a network of small-celled hyphae forms within
the epidermal wall, and haustoria are sent into the cell cavity
beneath. Ultimately each wall locule contains a single multinu-
cleate hyphal body, with a thin membrane. From this ascogenous
cell a single ascus arises as explained above. The intermediate
development is not clear, and it has not been possible to trace the

26 The University Science Bulletin

nuclear history from the rather scanty herbarium material at hand.
Cytological study of this fungus would be very desirable, since, if
the interpretations here given are correct, it is a highly aberrant
species, showing some resemblance to members of the genus Taph-
ridhim Lagerh. and Juel.

Although Nishida's paper is in Japanese, it may be assumed from
his English summary and from his figure that he missed the features
described above ('Vail habit" of mycelium, emergence of asci, pres-
ence of a pedicellate stalk cell).

15. Taphrina higginsii Mix
Taphrina higginsii Mix, Mycologia 39:71-76. 1947.

Causing small (up to 5 mm. diam.) yellowish unthickened spots
on leaves of Osmunda cinnamomea L.

Mycelium developing within outer epidermal wall, occasionally
forming unspecialized haustoria which project into the cell cavity.
A multinucleate hyphal element (ascogenous cell) forms in each
w^all locule, from which a single ascus emerges. Ascospores not
seen. Numerous spores (blastospores) appear in a peripheral layer
in the ascus before, during, or after emergence.

Asci hypophyllous or amphigenous, oblong-cyclindric, rounded at
apex, with a narrower cyclindric stalk cell, the lower part of which
is expanded into a foot within the wall locule. (Fig. 4, A.)

Dimensions: Of asci, 4:0-80^ X 13-23[jl; of stalk cells, 10-17[x X
8-lOix; of spores, 2-5[ji, X 1.5-4[jl.

Distribution: Georgia.

Material examined: Georgia: Hamilton, Pine Mountain, Blue
Springs Farm, May 26, 1938, B. B. Higgins.

This fungus, for which the writer is indebted to B. B. Higgins,
differs from Taphrina osmundae by its larger asci, and its narrow,
nearly cyclindric stalk cell. It also causes smaller spots, as is
natural on the more finely dissected fronds of the host. Like T.
osmundae it would repay cytological study.

An earlier collection made by W. A. Murrill at Blacksburg, Va.,
in 1897 could not be positively identified as T. higginsii. An ac-
count of this has been previously given (Mix, 1947).

16. Taphrina polystichi Mix
Taphrina polystichi Mix, Mycologia 30:563-579. 1938.

Causing small to large (up to 1 cm. diam.) j^ellowish or yellow-
brown thickened spots on leaves of Polystichum acrostichoides
(Michx.) Schott.

Mix: Monograph of Genus Taphrina 27

Mycelium intercellular.

Asci epiphyllous, occasionally also hypophylloiis, crowded, clav-
ate, rounded at apex, provided with a stalk cell, ascospores usually
eight, ovate to elliptic, sometimes budding in the ascus. (Fig. 4, C.)

Dimensions: Of asci, 23-53[jl X 4.5-8.5[x; of stalk cells, 13-23[ji. X
4-7[ji,; of spores, 3-6[x X 2-4[;..
- Distribution: eastern North America.

Material exaynined: New York: McLean, McLean Bog, July 5,
1940, A. J. M.; Oneida, June 10, 1935, H. D. House; Ringwood, iuly
23, 1940, A. J. M. North Carolina: Durham, 1935, F. A. Wolf;
Nantahala Gorge, May 29, 1941, A. J. M. Ohio: Hamilton Co.,
1934, Wm. Bridge Cooke. Pennsylvania: Lycoming Co., Trout
Run, June 6, 1925, L. 0. Overholts and L. T. Deniston (Herb. L. 0.
0. 9839). Tennessee: Cades Cove, May 30, 1937, L. R. Hesler.
West Virginia: Bayard, July, 1891, W. C. Sturgis.

This is the most frequently collected and apparently the most
common and widely distributed fern-inhabiting species of Taphrina
in, North America. It occurs throughout the eastern United States
and adjacent Canada. The first collection is that made by Sturgis
in W^est Virginia in 1891, and the first mention in literature is that
by Coker (1910) who called the fungus Exoascus filicinus.

17. Taphrina wettsteiniana Herzfeld

Taphrina wettsteiniana Herzfeld, Oesterreich Bot. Zeitschr. 60:249-254.
1910.

Causing small (up to 8 mm. diam.) roundish, well-defined, lemon-
yellow (brown in age) slightly thickened spots on under surfaces of
leaves of Polystichum lonchitis (L.") Roth. Spots scarcely visible
on the upper surface or only as a paler area.

Mycelium intercellular.

Asci hypophyllous or amphigenous, clavate, rounded at apex, with
or without a stalk cell; ascospores eight, fusiform or slightly curved,
usually with two globules. (Fig. 4, B.)

Dimensions: Of stalkless asci, 32-53pi X 6-lOiJ.; of stalked asci,
25-38[x X 6-lOiJ.; (17-23[/. X 5-7pL according to Herzfeld) ; of stalk
cells, 6-13[j. X 5-8[J.; of ascospores, 4.5-6.5[ji. X 2-4pi.. (Blastospores
may be elongate and bacterioid, 5 X 1[J!-)

Distribution: Tyrol.

Material examined: Tyrol: Sondestal ("sistendal des Geschniz-
tel, loc. class.") Aug., 1939, Fr. Wettstein.

Herzfeld, who described the fungus while working at Munich un-
der the direction of R. Wettstein, apparently designated no type

28 The University Science Bulletin

specimen. On inquiry at the Munich Museum it was learned that
preserved material of Herzfeld's collection was known to have been
in the museum but in 1938 it could not be found. However in 1939,
Dr. Fr. Wettstein kindly obtained fresh material from the place of
original collection and sent it to the writer. This material, de-
posited in the Mycological Herbarium, University of Kansas, is
chosen as the lectotype.

Although the mycelium is intercellular, very little modification
of the mesophyll occurs. The epidermis is browned.

18. Taphrina faulliana Mix
Taphrina faulliana Mix, Mycologia 30:563-579. 1938.

Causing small (up to 5 mm. diam.) round to oval, lemon-yellow
(brown in age or on drying) slightly thickened spots on leaves of
Polystichum, munitum (Kaulf.) Presl.

Mycelium subcuticular.

Asci hypophyllous, sometimes also epiphyllous, closely packed,
long-clavate, provided with a stalk cell. Ascospores not yet ob-
served, blastospores numerous, long-elliptic to bacilliform. (Fig. 4,
D.)

Dimensions : Of asci, 43-76[x X 6-9pL; of stalk cells, 13-33[jl X
4-7[jl; of blastospores, 4.5-6. 5[j. X 1.5-2[jl.

Distribution: Oregon, British Columbia.

Material examined: Oregon: Zigzag Mountain Trail, July 9,
1929, G. D. Darker (Arnold Arboretum, Path. Herb. 186) ; Rhodo-
dendron, Sept. 6, 1931, J. R. Hansbrough (Herb. J. R. H. 614).
British Columbia: Vancouver Island, Sidney or Lake Cowichan,
June, 1938, Malcolm Wilson.

In the original description of this fungus the spots are described
as brown. This must have been due to age of material since in the
freshly collected material received in 1938 from Malcolm Wilson,
the spots were, as he kindly pointed out, lemon-yellow in color. The
width of asci (4-7[ji.) given in the original description is in error.
Examination of the measurements recorded at the time of describing
shows the width to be 6-9pL as given above. Other revisions in di-
mensions (of stalk cells and conidia) result from study of additional
material.

The fungus was named in honor of Dr. J. H. Faull, from whom it
was first received. The first collection, apparently, was that of
Darker in 1929.

Mix: Monograph of Genus Taphrina

29

Fig. 4. Asci (X900) of A, Taphrina higginsii; B, T. wettsteiniana;
C, T . poly.-iUcItt ; D, T. fnulliana; E-H, T. tonduziana; E, F, ascogenous
cells in wall locules; G, emerging asci; H, asci.

30

The University Science Bulletin

19. Taphrina tonduziana P. Hennings
Taphrina tonduziana P. Hennings, Beibl. z. Hedwigia, 41:101-105. 1902.

Causing small (up to 5 mm. diam.) brown (with central areas
whitish by presence of asci) unthickened spots on leaves of Pteris
spinosa (L.) Desv.

Mycelium growing within the epidermal wall.

Fig. 5. A-E, Taphrina laurencia. A, B, mycelium in wall locule; C,
D, ascogenous cells; E, asci, in wall loculc; F, ascogenous cells (viewed
from above), G, asci of T. rhomboidalis. All X 900.

Mix: Monograph of Genus Taphrina 31

Asci amphigenous, emerging from wall lociiles, clavate, provided
with a stalk cell. Ascospores eight, long-elliptic to fusiform. (Fig.
4, E-H.)

Dimensions: Of asci, 20-33[jl X 7-8[;,; of stalk cells, 10-23tJ- X
4-7[jl; of spores, 4-7[jl X 2-3[x.

Distribution: Costa Rica.

Material examined: Costa Rica: San Jose, 1900, leg. A. Tondiiz,
comm. P. Hennings (Rehm, Ascomyceten 1461).

The errors in the original description by Hennings have already
been corrected and discussed (Mix, 1938), and the "wall habit" of
the fungus described.

20. Taphrina laurencia Giesenhagen
Taphrina laurencia Giesenhagen, Flora 76:130-156. 1892.

Causing remarkable, much branched, fine, bushy outgrowths on
fronds of Pteris quadnaurita Retz.

Mycelium growing within the epidermal wall.

Asci amphigenous, emerging from wall locules, provided with a
stalk cell. Ascospores oval to elliptic. (Fig. 5, A-E.)

Di7nensions: Of asci, 23-33[jl X 6-9[i; of stalk cells, 17-30pL X
4-7[ji,; of spores, 5-7pi. X 2-4[jl.

Distribution: Ceylon.

Material examined: Ceylon: Hakgala, March, 1914, T. Fetch;
Perideniya, 1912, T. Fetch (Ex. Herb. Rehm). India: G. Smith
(Ex. Herb. Sydow).

The correction of Giesenhagen's report that asci form within the
epidermal cells has been made earlier (Mix, 1938). Giesenhagen
did not find spores when describing the fungus but later (1901) re-
ported them to be in size near those of Taphrina vestergrenii.
Spores were present in the material examined by the writer.

21. Taphrina rhomboidalis Sydow and Butler
Taphrina rhomboidalis Sydow and Butler, Ann. Myc. 9:372-421. 1911.

Causing small yellow (becoming brown with age) rhomboidal
unthickened spots on leaves of Pteris quadriaurita Retz.

Mycelium growing within the epidermal wall.

Asci amphigenous, emerging from wall locules, provided with a
stalk cell; ascospores eight, ovate to elliptic. (Fig. 5, F, G.)

Dimensions: Of asci, 23-43[a X 8-10^.; of stalk cells, 23-43|x X
4-5pi,; of spores, 4-7[jl X 2-3. 5ii..

Distribution: Northern India.

Material examined: Himalaya: Kumaon, Barma Gori Valley,
June 24, 1907, Inayat Khan. Det. E. J. Butler. (Imp. Agric. Res.

32 The University Science Bulletin

Inst. I. 47.) Received from G. Watts Padwick, Imp. Agric. Res.
Inst. New Delhi, India.

The rhomboidal shape of the lesion is due to its being margined
by veins, though commonly more than one vein-islet is included.
The "wall habit" of the fungus, not fully described by the original
authors, has been discussed earlier (Mix, 1939) as well as the dif-
ferences between this species and the preceding. These are: the
production of unthickened spots instead of branched outgrowths, the
irregular, rather than polyhedral, outline of ascogenous cells when
viewed from above, and the possession by the ascus of a somewhat
narrower stalk cell.

22. Taphrina amplians Mix
Taphrina amplians Mix, Mycologia 31 :445-454. 1939.

Causing marked enlargement, but no thickening, of pinnae or
pinnules of Pteris orizabe Mat. and Gal. Affected areas becoming
golden-brown.

Mycelium growing within the epidennal wall.

Asci amphigenous, emerging from wall locules, golden-yellow
before maturity, and containing yellow material in epiplasm after
spore formation, provided with a stalk cell; ascospores eight, ovate
to elliptic. (Fig. 6. A, B.)

Dimensions: Of asci, 26-36[ji. X S-lOpL; of stalk cells, 23-40ix X
5-8[x; of spores, 5-6.5[jl X 2-3tJL.

Distribution: Guatemala.

Material examined: Guatemala: Quetzaltenango: Santa Maria
de Jesus, Dec. 28, 1936, J. H. Faull (Herb. J. H. F. 12939).

In studying material of Taphrina uleana (P. Henn.) Gies. the
writer was struck with the similarity of the healthy fronds of the
host to those of Pteris orizabe, and of the resemblance of the lesions
to those caused by T. amplians. This raised the question whether
the two hosts and perhaps the two fungi might be identical.

The ferns were submitted to C. A. Weatherby of the Gray Her-
barium, who compared them with authentic herbarium-specimens.
The host for Taphrina uleana is not Pteris decurrens Presl. as re-
ported by Hennings but Pteris defiexa Lk. Pteris defiexa and Pteris
orizabe are distinct, though closely related species, with different
geographical distribution.

The following specimens of Taphrina uleana were carefully
studied: 1. Museum botanicum Berolinense. E. Ule. Herbarium
Brasiliense No. 3076. "Taphrina (Exoascu^) Uleana P. Henn. Auf
Pteris decurrens, Estado de Rio de Janeiro, wald bei Tijuca, Marz.

Mix: Monograph of Genus Taphrina

33

Fig. 6. A, Asci, B, ascogenous cells (from above) of Taphrinn am-
AH ^'ofS' ^^^ °^ ^' ^"^^^^^"*/ E, asci of T. pteridis in wall locule.

3—3659

34 The University Science Bulletin

1900. leg. E. Ule. Typus!" 2. Six packets of : Mycotheca brasilien-
sis, No. 86, "Taphrina uleana P. Henn. in pterid. decurr. Rio de Jan.
Tijuca, 1900 E. Ule." Three of these packets were seen in the Far-
low Herbarium and three in the Botanical Museum, Stockholm.
One of the packets in the Farlow Herbarium is further designated,
"Ex. Museo. Botanico Berolinense. Original."

The herbarium-sheet bearing the packet of type material (1,
above) has affixed to it a portion of Henning's paper (1904) giving
a short Latin diagnosis, and a German description, translated as fol-
lows: "Whole leaves or large areas affected by the fungus, they are
at first thin membranous, yellowish, later thicken somewhat and
become stiff and dirty brown. Unfortunately could only find occa-
sional asci with ripe spores, therefore possible that size of asci may
be different: Not precluded that the species better placed in
Taphrina."

Most of this material is abominable. It gives the appearance of
having been wet and then dried without pressing. The type ma-
terial is the worst of all. In one of the packets in the Farlow Her-
barium the healthy portions of the fronds are well preserved and
it can be seen that the diseased areas are enlarged, but not thick-
ened, and browned. This, together with the description quoted
above, suggests that Taphrina amplians and T. uleana may be simi-
lar fungi.

Thorough study of all available material of T. uleana has failed
to reveal any mycelium or asci of Taphrina, any wall locules (even
empty) such as are produced by T. amplians, anything in fact, but
empty host cells and spores and mycelium of various molds. Study
of the type specimen was made before writing the description of T.
amplians; of the other specimens later. It seems posisble that the
writer may have described (as T. amplians) a fungus related to or
even identical with T. uleana. Since, however, the type specimen of
Taphrina uleana does not show any recognizable fungus, Ta'phrina
uleana must be declared a nomen dubium.

23. Taphrina ptendis Viegas
Taphrina ptendis Viegas, Bragantia 4:5-392. 1944.
Causing small (3-4 mm. diam.) round to elliptic (sometimes
angular) yellowish to brown, necrotic spots on leaves of Pteris sp.
Mycelium growing within the epidermal wall.
Asci amphigenous, clavate, rounded at apex, emerging from wall
locules; stalk cells cylindric; ascospores eight, elliptic. (Fig. 6, E.)
Dimensions: Of asci, 16-26[x; stalk cells, 8-20\i. X 6-7[jl; of asco-
spores, 4-7iJ. X 2.5-3[x.

Mix: Monograph of Genus Taphrina 35

Distnbiition: Campinas, Sao Paulo, Brazil.

Material examined: Brazil: Sao Paulo, Campinas, Bosque dos
Jequitibas, June 12, 1943, A. P. Viegas and R. 0. Botero (part of
type, received from Viegas).

The mycelium of this fungus is not subcuticular, as stated by
Viegas (1. c), but develops within the epidermal wall, the mature
asci protruding from a wall locule.

24. Taphrina lutescens Rostrup

Taphrina lutescens Rostrup, Meddel. Naturh. Foren. 1890. 246-264.
Magnusiclla lutescens (Rostr.) Sadebeck. Jahrb. Hamburg Wissensch. Anst.
10:5-110. 1893.

Causing small (up to 3 mm. diam.) pale yellow' unthickened spots
(the central portion becoming covered with a layer of asci), on
leaves of Thelypteris thelypteris (L.) Nieuwl.

Mycelium intercellular.

Asci hypophyllous, clavate, rounded at apex, lacking a stalk cell ;
ascospores eight, ovate to elliptic, commonly budding within the
ascus, filling it with bacterioid blastospores. (Fig. 6, C, D.)

Dimensions: Of asci, 27-75[jl X 5-11[jl (average 40[jl X 7[x) ; of as-
cospores, 3-6[xX2-4[ji, (average 4[ji,x2.6tJL) ; of blastospores, 1.5-3ijlX
1.5-2[ji.

Distribution: northeastern United States and adjacent Canada,
northern Europe.

Material examined: Maine: York, Aug., 1897, R. Thaxter.
New York: Shelter Island, Aug., 1901, Herb. W. G. Farlow. Wis-
consin: Madison, July 20, 1943, H. C. Green. Denmark: See-
land, Gjorslev, July, 1889, E. Rostrup; Lyngby, July, 1909 (received
from J. Dearness). Germany: Schleswig-Holstein, Pugum near
Glucksberg, July 13, 1908, 0. Jaap (Fungi sel. exsicc. 304) ; Zahlen-
dorf near Berlin, Sept., 1901, Sydow (Rehm, Ascomycetes 1413).
Latvia: Leinzale, Milskabe, Aug. 5, 1934, A. Kirulis (Fungi latvici
652).

HOST INDEX TO SPECIES OF TAPHRINA ON FERNS

Athyrium filix-jemina Roth.

Taphrina athyrii Siemaszko
Blechnum sp.

Taphrina blechni Bresadola ex Mix
Cystopteris fragilis (L.) Bernh.

Taphrina cystoptcridis Mix
Dryopteris arguta (Kaulf.) Wats.

Taphrina californica Mix
Dryopteris cheilanthoides (Kze.) C. Chr.

Taphrina ecuadorensis Syd.
Dryopteris filix-mas Schott.

Taphrina vestergrenii Gies.

36 The University Science Bulletin

Dryopteris lacera 0. Kuntze.

Taphrina vestergrenii Gies.
Dryopteris marginalis (L.) A. Gray.

Taphrina gracilis Mix
Dryopteris poiteana (Bory) Urban

Taphrina thaxteri Mix
Dryopteris rigida (Hoffm.) Underw.

Taphrina fusca Gies.
Dryopteris rigida var. australis Christ.

Taphrina jusca Gies.
Dryopteris spinulosa (Muell.) Kuntze

Taphrina athyrii Siemaszko

Taphrina filicina Rostr. ex Johans.

Taphrina jusca Gies.
Dryopteris sjnnulosa var. americana (Fisch.) Fern.

Taphrina filicina Rostr. ex Johans.

Taphrina jusca Gies.
Dryopteris spinulosa var. intermedia Underw.

Taphrina filicina Rostr. ex Johans.

Taphrina jusca Gies.
Dryopteris sp.

Taphrina jasciculata (Lagerh. and Sadeb.) Gies.
Onoclea sensibilis L.

Taphrina hiratsukae Nishida
Osmunda cinnamomea L.

Taphrina higginsii Mix
Osmunda regalis L.

Taphrina osmundae Nishida
Polystichum acrostichoides (Michx.) S'chott.

Taphrina polystichi Mix
Polystichum aristatum Presl.

Taphrina cornu-cervi Gies.
Polystichum lonchitis (L.) Roth.

Taphrina wettsteiniana Herzf.
Polystichum munitum (Kaulf.) Prsl.

Taphrina jaulUona Mix
Pteretis nodulosa (Michx.) Nieuwl.

Taphrina hiratsukae Nishida
Pteretis struthiopteris (L.) Todaro

Taphrina hiratsukae Nishida
Pteris orizabe Mat. and Gal.

Taphrina amplians Mix
Pteris quadriaurita Retz.

Taphrina laurencia Gies.

Taphrina rhomboidalis Syd. and Butl.
Pteris spinosa (L.) Desv.

Taphrina tonduziana P. Henn.
Pteris sp.

Taphrina pteridis Viegas
Thelypteris thelypteris (L.) Nieuwl.

Taphrina hiratsukae Nishida

Taphrina lutescens Rostr.

Mix: Monograph of Genus Taphrina 37

II. SPECIES ON SALICACEAE

Populus and Salix

25. Taphrina populina Fries

Taphrina populina Fries, Syst. Mycol. 3:520. 1832.

Taphrina aurea auct.

Exoascus populi Thiimen, Hedwigia 13 :97-98. 1874.

Exoasais aureus Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 1:93-124.

1884.

Exoascus flavo-aureus Cocconi, Mem. R. Accad. Sci. Inst. Bologna 4:187-

198. 1894.

Causing golden yellow, convex-concave spots, often small (5-10
mm. diam.), but sometimes confluent and large, involving half or
nearly all of leaf blade, on Populus angulata Ait., P. balsamifera
L., P. berolinensis Dipp., P. ciliata Wallr., P. generosa Henry, P.
laurifolia Ledb., P. nigra L., P. nigra L. var. italica Dur., P. petrow-
skyana Schneid., P. rasomowskyana Schneid.

Mycelium intercellular.

Asci variable in form, cylindric or clavate, rounded or truncate at
the apex, narrowed toward the base. Stalk cell present or absent,
variable in form, often triangular, wedge-shaped, or bluntly
rounded. Asci inserted between epidermal cells for as much as one-
third of their lengths. Asci when young with yellow oily contents,
large yellow globules persisting after spore formation. Ascospores
rarely seen; then commonly four, budding immediately so that
mature asci are filled with numerous blastospores. (Fig. 7, A-E,
Fig. 8, A-C.)

Dimensions: Of asci, 30-122pL X 13-30[ji.; of stalk cells (when
present) 4-27ijl X 8-23[jl; of ascospores, 4-6.5[ji,X4-5[ji.; of blasto-
spores, 2-3[JL X 2.5-1.5[a. Minute bacterioidal blastospores occa-
sionally seen.

Distribution: widely distributed in Europe, locally throughout
eastern North America, occasionally western North America, known
also from India, China, and Japan.

Material examined: Populus angulata. Germany: Brandenburg,
Spathsche Baumschulen zu Rixdorf b. Berlin, July 24, 1903, H. and
P. Sydow (Myc. Germ. 28).

P. balsamifera. ibid. (Myc. Germ. 28.) Norway: Akershus, As,
1885, F. Werenskiold.

P. berolinensis. Iowa: Ames, June 20, 1892, Pammel and Stewart
(as on P. " certinensis") . Norway: Akershus, Finsen in Aker (in
nursery), Sept. 10, 1931, I. J0rstad.

38

The University Science Bulletin

Fig. 7. Asci (X900) of Taphrina populina. A, on Populus anguluta;
B, P. balsamijera; C, P. berolinensis ; D, P. ciliata; E, P. generosa.

Mix: Monograph of Genus Taphrina 39

P. ciliata. India: Murree, July 20, 1908, E. J. Butler (Herb.
Crypt. Orient. Fungi) .

P. generosa. Norway: Akershus, As (in nursery) June 15, 1925,
I. J0rstad. Scotland: Murthly, July 15, 1925, J. S. Boyce (Herb.
J. S. B. 1565a) ; ibid. (Herb. J. S. B. 1565b) .

P. nigra. Iowa: Ames, June 20, 1892 (as on P. "betulifolia") .
Washington: Bremerton, July 20, 1912, E. Bartholomew (Fungi
Columb. 4488). West Virginia: Hillsboro, June 11, 1935, C. R.
Orton (as on P. "nigra var. babylonica") . Austria: between
Mossbrunnen and Marienthal, June, von Hohnel (Mus. Palat.
Vindob. Krypt. Exsicc. 1718). Italy: Venezia (Myc. Ital. 533).
Poland: Skiernewice, July 5, 1923, Z. Zweigboumawna. Scotland:
Murthly, July 15, 1925, J. S. Boyce (Herb. J. S. B. 1565c). Sweden:
Skane: Ystad, Bergsjoholmgard, July, 1939.

P. nigra var. italica. Iowa: Ames, June 20, 1892, L. H. Pammel
(as on "P. fastigiata") . Oregon: Portland, June 16, 1931, J. R.
Hansbrough (Herb. J. R. H. 610). Vermont: Duxbury, July 18,
1929, P. Spaulding (Herb. P. S. 66776). British Columbia:
Revelstoke, Sept. 7, 1930, J. R. Hansbrough (Herb. J. R. H. 610).
Quebec: Sillery, May 8, 1936, R. Pomerleau. Germany: Branden-
burg, Spathsche Baumschulen zu Rixdorf b. Berlin, July 24, 1903,
H. and P. Sydow (Myc. Germ. 30) .

P. petrowskyana. Iowa: Ames, June 20, 1892, F. C. Stewart.

P. rasomowskyana. Germany: Dahlem b. Berlin, July, 1902, P.
Hennings (Rehm, Ascomyceten 273b) .

The occurrence of a stalk cell in Taphrina populina was first re-
ported by Frank (1880) and later by Johanson (1887). The ex-
istence of a stalk cell was denied by Sadebeck (1890) , but as pointed
out by Johanson, Sadebeck had confused this fungus with Taphrina
rhizophora. Sadebeck later (1893) distinguished two types of asci
in T. populina: (1) Slender, deeply inserted asci, 80-112[jl X 20-27[j,,
provided with stalk cells and occurring on Populus nigra L. var.
italica Dur.; and (2), broad asci, 80[Ji. X 30-40pL, slightly inserted,
often lacking stalk cells, occurring on Populus nigra L. The fact
that the stalk cell may be present or absent has been recorded by
most later investigators.

In these studies the two types of asci mentioned by Sadebeck have
been found on all but one of the hosts listed above. Asci without
stalk cells may have a bluntly rounded or wedge-shaped base, and
are less deeply inserted than the longer asci with stalk cells. Neither
type of ascus occurs exclusively on any one host. In particular the
limits of ascus-size observed for Populus nigra are: 30-122[jl X

40

The University Science Bulletin

Fig. 8. Asci (x900) of Taphrina populina. A, on Populus nigra; B,
P. nigra var. italica; C, P. rasomowskyana.

Ascus Presence

Stalk cell

dimensions of stalk

dimensions

in microns cell

in microns

33-76 X 20-27 +

10-23 X 10-20

63-116X17-26 +or —

8-20 X 10-20

43-80 X 17-26 + or —

8-17 X 10-22

73-122 X 23-33 + or —

7-10 X 17-23

56-76 X 16-20 + or —

8-10 X 10-13

33-83 X 16-26 + or —

8-20 X 10-16

30-122 X 15-30 + or —

5-26 X 10-20

Mix: Monograph of Genus Taphrina 41

16-30[ji,, and for P. nigra var. italica: 42-109[jl X 15-30[jl. (More
specimens of the former have been studied.) Asci with stalk cells
seem equally common on both hosts. Incidentally no asci as wide
as those reported by Sadebeck have been observed.

A further example of the variability of Taphrina populina appears
in the attempt by Cocconi (1894) to erect a new species, Exoascus
fiavo-aureus for the fungus on P. nigra var. italica. It possesses,
according to his observations, a tapering, rhizoidal ascus-base, in-
serted between the epidermal cells of the host.

The following is a record of the writer's studies of Taphrina
populina on the various host-species listed above:

Host
Populus angulata Ait.
Populus balsamifera L.
Populus berolinensis Dipp.
Populus ciliata Wallr.
Populus generosa Henry
Populus laurijolia Ledeb.
Populus nigra L.
Populus nigra L. var. 40-lOQ X 15-30 + or — 10-30 X 10-20

italica Dur.
Populus petrowskyana Schneid. 43-76 X 13-26 -|- or— 10-23 X 13-20

Populus rasomowskyana Schneid. 35-86 X 17-30 4- or — 7-30 X 10-20

Choosing the fungus on P. nigra as the type, it might be possible
to distinguish varieties, one for each host-species. The danger in
this would be that if additional specimens were studied some of the
differences evident here might disappear. In the absence of infor-
mation as to the biological specialization it seems best merely to
record that Taphrina populina shows marked variability on different
hosts.

26. Taphrina populi-salicis Mix
Taphrina populi-salicis Mix, Trans. Kansas Acad. Sci. 50:77-83. 1947.

Causing golden-yellow convex-concave spots on leaves of Populus
fremontii S. Wats., P. trichocarpa Torr. and Gray, and Salix lae-
vigata Bebb.

Mycelium intercellular.

Asd hypophyllous, oblong-cylindric, rounded at the apex. Stalk
cell invariably present, short and wedge-shaped or long and taper-
ing, often forked, deeply inserted in leaf tissues. Spores (blasto-
spores) numerous, elliptical, ovate, or narrowly elongate. (Fig. 9,
A, C, Fig. 10, A, B.)

Dimensions: Of asci, 50-106[jl X 13-30[x; of stalk cells, 7-92[jl X
7-27[jl; of spores, 1.5-5|a X 0.5-4.5[i.

Distribution: Pacific Coast States.

42

The University Science Bulletin

Fig. 9. Asci (X 900) of Taphrina po-puli-salicis. A, on Populics }re-
montii; B, C, P. trichocarpa.

Mix: Monograph of Genus Taphrina 43

Material examined: P. fremontii. California: Berkeley, July
10, 1938, H. N. Hansen; Palo Alto, San Francisquito Creek, May
13, 1941, R. H. Thompson; Sacramento, June 8, 1924, H. E. Parks
and W. S. Fields.

P. trichocarpa. California: Glendale, May, 1897, A. J. Mc-
Clatchie (Fungi Columb. 1228) ; Palo Alto, April 17, 1940, R. H.
Thompson; Richardson Grove, Aug. 22, 1929, G. D. Darker. Ore-
gon: Lane Co., Eula, June 13, 1920, J. S. Boyce (Herb. J. S. B.
594) ; Portland, Aug. 12, 1931, J. R. Hansbrough (Herb. J. R. H.
609). British Columbia: New Westminster, July, 1938, Malcolm
Wilson; Owl Creek, June 24, 1931, J. R. Hansbrough and T. S.
Buchanan (Herb. J. R. H. 608).

Salix laevigata. California: Lassen Co., Mineral, Sept. 10,
1911, E. P. Meinecke (Herb. J. S. B. 282).

This fungus, while causing leafspots quite like those caused by
Taphrina populina shows definite morphological differences, and an
apparent biologic distinction, from that species. Boyce (1927) re-
ports observing in the Pacific northwest that Populus nigra often is
not attacked in the same locality in which P. trichocarpa is affected,
and that in a nurseiy in Scotland, in 1925, Taphrina populina at-
tacked Populus generosa, P. laurifolia, and P. nigra (the last named
being the most heavily infested), while P. trichocarpa growing with
its leaves intermingling with those of the diseased P. generosa re-
mained healthy.

It is of interest that indigenous west coast species of Taphrina
described by Mix (1939) and Ray (1939) are different from related
forms found in the eastern United States. {Taphrina populina was
undoubtedly introduced into this region along with its host.)

27. Taphrina johansonii Sadebeck

Taphrina johansonii Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 8:61-95.

1890.

Taphrina rhizophora Johanson, Bih. t.k. Svensk Akad. Handl. 13:3-28.

1887, in part.

Exoascics johansonii Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 10:5-110.

1893.

Ezoascus aureus auct.

Causing golden-yellow enlargements of carpels of Populus canes-
cens Reichenb., P. grandidentata Wats., P. sieboldii Miq., P. tremula
L., P. tremula var. pendula Loud., P. tremuloides Michx.

Mycelium intercellular.

Asci elongate, clavate, rounded at apex, with no stalk cell but with
a tapering base deeply inserted between host cells ; when young with

44

The University Science Bulletin

fjG. 10. Taphrina populi-salicis. A, ascogenous cells on Populus tricho-
carpa; B, asci on Salix laevigata. X 900-

Mix: Monograph of Genus Taphrina 45

yellow oily contents, yellow oil globules persisting after spore for-
mation; ascospores rarely seen, budding at once to fill the ascus with
numerous blastospores which are variable in form: oval, fusiform,
or rod-shaped. (Fig. 11, A-E.)

Dimensions: Of asci, 60-145[jt. X 12-27[jl, narrowing at the foot to
a width of 3 to 10[x, inserted for about one-third (occasionally one-
half) the ascus-length ; of ascospores, 4-7[ji. X 4-6[jl; of blastospores,
4-lOpL X 1.5-41J..

Distribution: Europe, eastern North America, Japan.

Material examined: P. canescens. Russia: Moscow, 0. L. Park,
June, 1893, P. Sydow (Myc. March. 3820).

P. grandidentata. Massachusetts: Peabody, April 19, 1938, D.
H. Linder.

P. sieboldii. Japan: Morioka, Kaminoyai, May 24, 1908 (Herb.
Morioka Imp. Coll. Agric. and For.).

P. tremida. Denmark: Jylland, Viborg, May 28, 1890, C. A.
Gad (Fungi Scand., Gunnar V. Schotte) ; Fireso, Sj. Koningaarden,
May 6, 1894, F. K0lpin Ravn. Finland: Helsingfors, May, 1903,
J. I. Liro (Vestergren, Micr. Rar. Sel. 1619).

P. tremula var. pendula. Sweden: Stockholm, Hortus Bergianus,
June 7, 1924, T. Vestergren.

P. tremuloides. Massachusetts: Medford, April 25, 1891, A.
B. Seymour and Agnes W. Lincoln (Seymour and Earle, Econ. F.
191 as T. rhizophora) ; Wellesley, May 20, 1935, D. H. Linder.
Michigan: between Brighton and Whitmore Lake, May 24, 1935,
L. E. Wehmeyer; near Mich. Agric. Coll., May, 1892 (N. A. F.
1885, F. Columb, 1312, as T. rhizophora). New York: Ithaca,
KHne Woods Road, May 22, 1940, A. J. M.; Connecticut Hill, May
28, 1940, P. A. Readio. Wisconsin: Racine, F. L. Stevens (F.
Columb. 407, as T. rhizophora) .

Species on the carpels of Poipulus were not at first distinguished
from Taphrina populina [T. aurea) . Thus Farlow (1878) reported
T. aurea on carpels of P. grandidentata. Following the description
of T. rhizophora on carpels of P. alba by Johanson (1887) American
authors accepted that name.

Farlow (1888) lists as occurring in North America: T. rhizophora
on carpels of P. fremontii, P. grandidentata, "P. pyramidalis," and
P. tremuloides. The record on P. fremontii seems to be an error,
referring to a collection of T. populi-salicis on leaves of that host. "P.
pyramidalis" is a varietyof P. alba, but the writer has been unable
to find any record of such a collection made in North America.

46

The University Science Bulletin

P"'iG. 11. Taphrina johansonii. Asci (X900), A, on Populus tremula;
H, /■'. canesccris ; C, P. sieboldii; D, P. tremuloides ; E, P. grandidentata.

Mix: Monograph of Genus Taphrina 47

Farlow's fungus should have been reported as Taphrina johan-
sonii. This was pointed out by Patterson (1895) following the
description of T. johansonii by Sadebeck (1890). Sadebeck, who
earlier (1884) had called this fungus Exoascus aureus, described it
as having asci 92-105ij, long, inserted but never showing rhizoidal
forking, it being distinguished by these characters from T. rhizo-
phora whose asci were 120-160[ji. long and often showed rhizoidal
extensions of the ascus-base.

The studies reported here, although they extend the limits of
ascus-size beyond those reported by Sadebeck, reveal no significant
differences between the forms on the various hosts listed above. The
ascus-dimensions observed are: For P. canescens, 60-125[x X 17-26[x;
for P. grandidentata, 66-122[x X 13-23[ji.; for P. sieboldii, 6O-I2O1;, X
17-26[jl; for P. tremida, 60-145[Ji X 17-27[J(,; for P. tremuloides,
60-125[jL X 16-26[JL. All these host-forms show asci deeply inserted
and possessing a rhizoidal foot, occasionally branched. Clearly
they are all Taphrina johansonii.

Ikeno (1901) studying a form on Populus tremula var. villosa
Wesm. considered it to be intermediate between T. johansonii and
T. rhizophora, its asci measuring 83-133pi, X 20-27[x and penetrating
the host for from 17-20[;l. He did not observe any rhizoidal exten-
sions and on this basis decided to call it T. johansonii. Its dimen-
sions fall within the limits of ascus-size for T. johansonii on P.
tremula as given above.

28. Taphrina rhizophora Johanson

Taphrina rhizophora Johanson, Bih. t. K. Svensk. Vetensk. Akad. Handl.

13:3-28. 1887.

Taphrina aurea auct. in part.

Exoascus aureiis auct. in part.

Exoascus rhizophorus (Johans.) Sadebeck, Jahrb. Hamburg. Wissensch.

Anst. 10:5-110. 1893.

Causing golden-yellow enlargements of carpels of Populus alba L.

Mycelium intercellular.

Asci long, clavate, rounded at apex, attentuate at the base to a
long, rhizoidal, often forking end, deeply inserted (as much as half
the ascus-length) between host-cells. Young asci with yellow con-
tents; yellow globules persisting after spore-formation. Ascospores
rarely seen, budding at once to form numerous small blastospores,
filling the ascus. (Fig. 12, A).

Dimensions: Of asci, 76-198[jl X 20-30[x, tapering portion 2-lOpL
in diam.; ascospores reported by various authors as round with a
diameter of 4[jl; blastospores, 3.5-5\t. X 3-4^^.

48

The University Science Bulletin

Fig. 12. Asci (X 900) of A, Taphrina rhizophora; B, T. coryli.

Mix: Monograph of Genus Taphrina 49

Distribution: Europe.

Material examined: Populus alba. Algier: Teniet-el-Had,
April 20, 1930, R. Maire (Herb. R. M., Champ. Afr. d. Nord. 9933,
on P. alba var "hickeliana") . Poland: Pulawy, May 4, 1930, K.
Jankowska (Herb. Inst. Phytopath. Schol. Sup. Agric. Warsaw).
Sweden: Skane, Alnarp, May 19, 1890, F. Ulrichsen (Ericksson, F.
Par. Scand. 358, as on Populus sp. but probably on P. alba).

The first mention of a form of Taphrina on the carpels of Populus
alba and of P. tremula is by Magnus (1874) who called it Taphrina
aurea. Sadebeck (1884) considered the fungi on leaves and on
carpels of Populus to be the same, calling them Exoascu^ aureus.
Johanson (1887) described the form occurring on carpels of Populus
alba and P. tremula as a new species, Taphrina rhizophora. He gave
dimensions of the asci as 80-156pi. X 16-22[jt,, and described the deep
insertion of the asci and their long, rhizoidal, often forked, basal
extensions. (He suggested that the fungus known in North America
on P. tremuloides was the same.) Sadebeck (1890) in recognizing
T. rhizophora, separated the form on P. tremula as a new species,
Taphrina johansonii. He gave the ascus-length of T. rhizophora
as 120-160pL, showing no overlapping with T. johansonii, and these
measurements have been copied in subsequent literature. Actually
there is an overlapping, the writer's measurements (above) showing
asci of T. rhizophora as short as 76tJL.

The writer's stay in Sweden in 1939 was interrupted before a
visit to Uppsala could be made, and Johanson's type material has
not been studied. This does not seem important since the larger
size of the asci on P. alba is quite apparent.

Following the principle of disturbing existing situations as little
as possible both T. rhizophora and T. johansonii are here recognized
as valid species, though it is not clear that they are well distin-
guished. Branching rhizoidal extensions of the asci are as common
in the one fungus as in the other, and there is an overlapping in
ascus-size between the two. Consistent treatment of species on
Populus would call for uniting these two species into one, or for di-
vision of Taphrina populina into several species.

The occurrence of T. rhizophora on P. tremula reported by the
writer (1936a) was an error, due to failure to recognize that Sade-
beck meant to separate the fungus on P. tremula as T. johansonii.
Jaczewski (1926) reports T. rhizophora as occurring on Populus
bachofeni. This host, according to Index Kewensis, is identical with
P. alba.

4—3659

50 The University Science Bulletin

HOST INDEX TO SPECIES OF TAPHRINA
ON SALICACEAE

Pcypulus alba L.

Taphrina rhizophora Johans.
Populus angulata Ait.

Taphrina populina Fr.
Populus balsamifera L.

Taphrina populina Fr.
Populus berolinensis L.

Taphrina populina Fr.
Populus canescens Reichenb.

Taphrina johansonii Sadeb.
Populus ciliata Wallr.

Taphrina populina Fr.
Populus fremontii S. Wats.

Taphrina populi-salicis Mix
Populus generosa Henry

Taphrina populina Fr.
Populus grandidcnta Wats.

Taphrina johansonii Sadeb.
Populus laurifolia Ledeb.

Taphrina populina Fr.
Populus nigra L.

Taphiina populina Fr.
Populus nigra L. var. italica Dur.

Taphrina populina Fr.
Populus petrowskyana Schneid.

Taphrina populina Fr.
Populus rasomoioskyana Schneid.

Taphrina populina Fr.
Populus sieboldii Miq.

Taphrina johansonii Sadeb.
Populus tremula L.

Taphrina johansonii Sadeb.
Populus tremula L. var. pendula Loud.

Taphrina johansonii Sadeb.
Populus tremula L. var. villosa Wesm.

Taphrina johansonii Sadeb.
Populus tremtdoides Michx.

Taphrina johansonii Sadeb.
Populus trichocarpa Torr. and Gray

Taphrina populi-salicis Mix
Salix laevigata Bebb

Taphrina populi-salicis Mix

Mix: Monograph of Genus Taphrina 51

III. SPECIES ON BETULACEAE

Corylvs, Ostrya, Carpinus, Betula, Alnus
29. Taphrina coryli Nishida
Taphrina coryli Nishida, Miyabe Festschrift, Tokyo, 1911, pp. 157-212.
Causing leaf curl involving small spots, large areas, or the whole
blade, of Corylus americana Marsh., C. heterophylla Fisch., C. ros-
irata Ait., and C. sieboldiana Blume. Affected areas at first yellow-
ish, becoming white at maturity of asci.
Mycelium intercellular.

Asci amphigenous, cylindric or cylindric-clavate, rounded or trun-
cate at the apex, seated on or slightly inserted in a stalk cell which
may be a little broader than the ascus. Ascospores eight, globose,
ovate, or elliptic, often budding in the ascus. (Fig. 12, B.)

Dimensions: Of asci, 20-40tJL X 8-12ijl; of stalk cells, 10-20[ji. X
8-n\).; of spores, 4-6.5[jl X 3.5-6[a.

Distribution: eastern North America, Japan.

Material examined: Corylus americana. Connecticut: Sandy
Hook, Botsford Hill, July 4, 1928, A. J. M.; Hammertown Road, be-
tween Botsford and Monroe, July 20, 1947, id. Massachusetts:
Andover, Harold Parker Forest, June 3, 1939, D. H. Linder. Wis-
consin: Madison, June 2, 1915, J. J. Davis (F. Columb. 4882);
Sparta, June, 1916, J. J. Davis.

C. heterophylla. Japan: Iwate, Terada, June 8, 1907, K. Sa-
wada (Herb. Morioka Imp. Coll. Agric. and For.).

C. rostrata. Connecticut: Westville, summer 1890, R. Thaxter
(Rel. FarL642).

C. sieboldiana. Japan: Iwate, Mt. Himekawe, June 30, 1907, K.
Sawada (Herb. Morioka Imp. Coll. Agric. and For.).

30. Taphrina ostryae Massalongo

Taphrina ostryae Massalongo, Bot. Centbl. 34:389-390. 1888.

Causing small, brown, definitely margined, unthickened spots on
leaves of Ostrya carpinifolia Scop.

Mycelium subcuticular.

Asci hypophyllous, oblong, rounded or obtuse at the apex, seated
on a stalk cell which may be somewhat wider than the ascus. As-
cospores eight, globose. (Fig. 13, A.)

52 The University Science Bulletin

Dimensions: Of asci, 20-24[jl X 12-14pL (Massalongo) , 17-26ijl X
T-lOpL (Mix); of stalk cells, variable size (Massalongo), T-IO^jl X
7-13[K (Mix) ; of spores, 4.5-5tJi. X 4-4.5tJi,.

Distribution: Italy.

Material examined: Italy: Lucca, Vellana, Oct., 1890, G. Tog-
nini (Br. and Cav. F. Par. D. PI. Colt, od Ut. 169) ; Verona, valley
Tregagno, July, 1898, C. Massalongo (loc. class.).

31. Taphrina virginica Sadebeck

Taphrina virginica Sadebeck, Ber. Deutsch. Bot. Ges. 13 :265-280. 1895.

Causing small, pale yellowish thickened areas or leaf curl of the
whole blade, on leaves of Ostrya virginica Willd.

Mycelium intercellular.

Asci epiphyllous, sometimes hypophyllous, rounded or truncate at
the apex, with rounded or truncate, sometimes broadened base, lack-
ing a stalk cell. Ascospores eight, ovate to elliptic, often budding in
the ascus. (Fig. 13, B.)

Dimensions: Of asci, 20-46[jl X 8-13ijl, the widened base occa-
sionally measuring ITpi.; of spores, 4-6. Sj;, X 4-5ijl.
Distribution: eastern North America.

Material examined: Kansas: Vinland, Baldwin Woods, May 15,
1947, A. J. M.; Lecompton, May 15, 1947, R. L. McGregor. Massa-
chusetts: Princeton, July, 1888, W. G. Farlow; Wellesley, June 8,
1888, A. B. Seymour (E. F. 497a. Duplicate of type) . New Hamp-
shire: Temple, June 16 and 18, 1888, A. B. and A. C. Seymour (E.
F. 497b, and 497c). Ve:rmont: Ferrisburg, Shellhouse Mtn., July
23, 1884, Faxon. Wisconsin: Potosi, June 7, 1918, J. J. Davis.
Ontario: Woods east of Holland River, June 3, 1937, H. S. Jackson
(Crypt. Herb. Univ. Toronto, 12066).

32. Taphrina carpini (Rostr.) Johanson

Taphrina carpini (Rostr.) Johanson, Ofv. Kongl. Svensk. Vetensk. Akad.

Forhandl. 1885:29-47. 1886.

Exoascus carpini Rostrup, Bot. Centbl. 5:153-154. 1886.

Causing witches' brooms of Carpinus betulus L. and C. orientalis
Mill.

Mycelium subcuticular.

Asci hypophyllous, cylindric, rounded at the apex, truncate and
sometimes broadened at the base, lacking a stalk cell; ascospores
eight, ovate to elliptic. (Fig. 13, C.)

Dimensions: Of asd, 20-30ijl X 7-14[jl (may broaden at base to
20-2411.) ; of spores, S.5-5[l X 3-4.5tJL.

Mix: Monograph of Genus Taphrina

53

Fig. 13^^ Asci (X 900) of A, Taphrina ostryae; B, T. virginica; C, T.
carpim; D, T. australis; E, T. betulicola; F-I, T. americana' F G on
Betula jontmahs; H, on B. papyrijera; I, on B. lutea. ' '

54 The University Science Bulletin

Material examined: Carpinus betulus. England: Surrey, Vir-
ginia Water, July 22, 1935, A. J. M. Germany: Thuringia, Steiger
near Erfurt, May 29, 1920, H. Diedicke; Escheberg near Bergsdorf,
June 20, 1909, 0. Jaap (F. Sel. Exs. 408).

C. onentalis. Russia: Kubinski, Ascobaidjan, May 30, 1927;
Uljanishtshev (Herb. Inst. Prot. Plants, Sect. Phytopath., Lenin-
grad). Sweden: Lund, June 11, 1887, E. Ljungstrom (Fungi Sue-
cici).

33. Taphrina australis (Atk.) Geisenhagen

Taphrina australis (Atk.) Giesenhagen, Flora 81:267-361. 1895.
Exoascus australis Atkinson, Bull. Torr. Bot. Club 21:372-380. 1894.

Causing leaf-curl (without evident thickening) of small areas or
of the whole blade of leaves of Carpinus caroliniana Walt.

Mycelium subcuticular.

Asci epiphyllous, cylindric, truncate or rounded at the apex,
narrowed or somewhat broadened at the base, lacking a stalk cell.
Ascospores eight, ovate to elliptic. (Fig. 13, D.)

Dimensions: Of asci, 20-53t;. X 7-12[j, (asci as long as 60[j(. re-
ported by Atkinson) ; of spores, 4-6a X 2.5-4.5[x.

Distribution: eastern North America.

Material examined: Alabama: Auburn, April 30, 1892, G. F.
Atkinson (Type? Herb. Dept. Plant Path. Cornell Univ. Atkinson
Coll. 2401). Connecticut: North Bloomfield, May, 1919, P.
Spaulding (Herb. U. S. D. A. Div. For. Path. 45975).

This species seems to be well distinguished from the preceding
by its larger asci and by not forming witches' brooms.

34. Taphrina betulicola Nishida

Taphrina betulicola Nishida, Miyabe Festschrift. Tokyo. 1911.

Causing witches' brooms on Betida ermani Cham.

Mycelium subcuticular, perennial.

Asci cylindric, rounded or truncate at the apex, provided with a
cylindric stalk cell. Ascospores eight, ovate to elliptic, frequently
budding in the ascus. (Fig. 13, E.)

Dimensions: Of asci, 26-46i;, X 13-22[jl; of stalk cells, 7-13(1 X
13-20^^; of spores, 3.5pi. X 3tJL. (Nishida's measurements of stalk cells
were: 8-20ijl X 20-26[i.)

Distribution: Japan.

Material examined: Betula ermani. Japan: Nikko, Mt. Shiraue,
July, 1907, S. Kusano. (Apparently part of type collection. Re-
ceived from Kusano).

Mix: Monograph of Genus Taphrina

DO

Fig. 14. A-E, asci of Taphrina hetulina. A, on Beiula carpatica; B,
B. intermedia; C, B. pubescens; D, B. pubescens (as T. lagerheimii) ;
E, B. pubescens (as T. lapponica) ; F, asci of T. splendens. All X 900.

56 The University Science Bulletin

This species is, for the present, held to be distinct from T. betulina.
Though its ascus-dimensions fall within the size-range of that spe-
cies, the average size (35 X 16^;.) is less, and the stalk cell is shorter.

35. Taphrina betulina Rostrup

Taphrina betidina Rostrup, Tidsskr. f. Skovbrug. 6:199-300. 1883.
Exoascus betulinus (Rostr.) Sadebeck, Jahrb. Hamburg. Wissensch. Anst.

10:5-110. 1893.

Taphrina lagerheimii Falm, Arkw.Bot. 15 -.1-41. 1917.

Taphnna lapponica Juel, Svensk. Bot. Tidsskr. 6:353-372. 1912.

Exoascus lapponicus (Juel) Jaczewski, Pocket Key for determination of

fungi. Part I. Exoascales, Leningrad, 1926.

Taphrina turgida (Sadeb.) Giesenhagen, Flora 81:267-361. 1895.

Exoascm turgidus Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 1:93-124.

1884.

Taphrina willeana Svendsen, Nyt Mag. Vidensk. 40:363-368. 1902.

Causing witches' brooms. In early stages several or all of the
leaves of a shoot may be yellowed (sometimes slightly enlarged but
not thickened) , but the adventitious twigs of a typical broom may
not be present. Subsequently a witches' broom develops. Oc-
curring on: Betula aurata Bechst., B. carpatica Waldst., B. inter-
media Thomas, B. nana L., B. pendula Roth., B. pubescens Ehrh.

Mycelium subcuticular, perennial.

Asci cylindric, rounded or truncate at the apex, provided with a
stalk cell which may be broad and seated, or wedge shaped and
inserted between epidermal cells. Ascopores eight, ovate to elliptic,
frequently budding in the ascus, filling it with smaller, ovate or
elliptic blastopores. (Fig. 14, A-E, Fig. 15, A,B.)

Dimensions: Of asci, 23-73ii X 10-26[jl; of stalk cells, 7-21[x X
10-30[;.; of ascospores, 4.5-6. 5[x X 4-5. 5[jl; of blastospores, 3.5-G[x X
2-4.5^;..

Distribution: Europe.

Material examined: Betida aurata. Germany: Hamburg, Ep-
pendorfer Moor, June 19, 1906, 0. Jaap (F. Sel. Exs. 352).

B. carpatica. Germany: Scheibe near Schandau, June, 1909, W.
Krieger (F. Sax. 2054).

B. intermedia. Norway: Gran Hadeland, top of Mt. Framstad-
saeterfjeld, June 18, 1901, N. Wille (type of T. willeana Svends.,
received from I. J0rstad). Sweden: Jiimtland, Storlien, Aug. 24,
1939, A. J. M. (two collections) ; Lappland, Abisko, Aug. 17-21,
1939, A. J. M. (ten collections) ; Palnoviken, Aug. 20, 1939, A. J.
M.; Kaskajaure, July 15, 1938, Th. Arwidsson (as T. lapponica).

B. nana. Norway: Finnmark, Bossekop in Alta, July 29, 1924,
I. J0rstad.

Mix: Monograph of Genus Taphrina

57

Fig. 15. A, B, asci of Taphrina beiulina; A, on B. peiidula {T. lur-
gida) ; B, B. intermedia {T. willeana) ; C, asci of T. splendcus on B.
pubescens. All X 900.

58 The University Science Bulletin

B. pendula (as T. turgida). Germany: Brandenburg, Bredower
Forest near Nauen, June, 1904, H. Sydow (Myc. Germ. 249).
Sweden: Grisslehamn, July 13, 1912, 0. Juel; Stockholm, Haga,
June 12, 1910, B. Palm, Norra Djurgarden, June 30, 1939, A. J. M.;
Tungelsta, July 13, 1939, A. J. M.

B. pubescens. Germany: Westphalia, Hauberg near Siegen,
May 22, 1920, H. Ludwig (Myc. Germ. 1646). Ireland: County
Wicklow, Killadreenan House, June 23, 1935, A. J. M. Sweden:
Jamtland, July 13, 1931, A. G. Eliasson; Storlien, Skurdalshojden,
July 15, 1936, T. Arwidsson; Storlien, Aug. 24, 1939, A. J. M. (two
collections); Kalama, B. Palm; Quikjokk, 1887, G. Lagerheim
(Roum. F. G. 4561).

B. pubescens (as T. lapponica). Norway: Troms, Kirkenesmoen
in Malselv, July 31, 1926, I. J0rstad. Sweden: Harjedalen,
Fjallnas, Malmagsvalen, June 28, 1933, A. G. Eliasson; ibid., July
2, 1933, A. G. E. ; Jamtland, Frostviken, Jorniklumpen, July 2, 1934,
J. A. Nannfeldt (F. Exs. Suec. 651) ; Lappland, Bjorkliden, July
19, 1911, 0. Juel (type) ; Abisko, July, 1907, T. Vestergren (Micr.
Rar. Sel. 1620. "Vidit auctor") .

B. pubescens (as T. lagerheimii) . Sweden: Lappland, Abisko,
Vaggejokk, Vilkisorta, Bjorkliden, Palnoviken, B. Palm (several
undated collections).

Synonymy : a. Taphrina turgida (Sadeb.) Gies.

Rostrup (1883) described Taphrina betidina from "birch" with-
out giving the* specific name of the host, though later (1890) he
stated that it was Betula odorata {B. pubescens). Sadebeck (1884)
described Exoascus turgidus, also from "birch," and gave Taphrina
betulina as a synonym. Later Sadebeck (1893) reported the host
for T. turgida as B. verrucosa {B. pendula) , and stated that T.
betulina occurred on B. pubescens. Sadebeck's distinction between
the two species was based on the form of the stalk cell, T. betulina
having a broad stalk cell, truncate or rounded at the base, seated on
the epidermis, while T. turgida had a wedge shaped stalk cell,
pointed below, and inserted between the epidermal cells. (He also
stated that the two species differed in the manner of formation of
their ascogenous cells.)

Rostrup (1890) expressed the opinion that Taphrina betulina and
T. turgida were identical, but later (1896) he agreed with Sade-
beck's distinction between the stalk cells of the two forms. He
also suggested a biological distinction, since he had observed Betula

Mix: Monograph of Genus Taphrina

59

pubescens and B. pendula growing close together, with only the
former showing witches' brooms.

Actually there is no morphological distinction between the fungus
on Betula pubescens and the one on B. pendula, either in size and
shape of asci or in form of stalk cells. In all specimens examined
from the hosts named above, both the broad, flat type of stalk cell,
and the wedge-shaped, inserted one have been seen. Sadebeck
himself (1893) stated that the stalk cells of T. turgida were very
variable, and that it would be difficult to tell the two species apart
by means of this character.

b. Taphrina lapponica Juel.

Juel (1. c.) in distinguishing T. lapponica from T. betuUna stated
that the former did not form witches' brooms though invading
"whole shoots." According to his account all of the leaves on a
branch became yellowed and sometimes enlarged, but adventitious
buds were not formed. In the summer of 1939, during a stay in
Lappland and Jamtland, the writer become convinced that the
above-described appearance is that of a young stage of a witches'
broom, resulting from infections of the current year or of a recent
year. Stages intermediate between "whole-shoot infection" and
witches' brooms were common.

Juel did not find any marked difference in ascus-size between the
two species, but reported ascus-dimensions for T. lapponica as
40tJL X 16[J., the dimensions for T. betulina being 45-55[;, X 15-20[j,.
He did, however, assign a shorter host list, finding T. lapponica to
occur on Betula intermedia and on B. pubescens.

That the two forms cannot be distinguished morphologically may
be seen from the following tabulation made mostly from specimens
collected by the writer in Sweden in 1939.

Taphrina lapponica, not forming witches' brooms.

Ascus-size
in microns

Stalk cell -size
in microns

45-50 X
30-50 X
40-73 X
23-50 X
30-50 X
33-66 X
36-50 X
36-50 X
36-60 X
33-66 X
30-46 X
26-43 X
33-50 X

17-23
13-20
17-20
13-20
13-17
13-18
13-20
13-20
17-20
13-20
12-20
17-20
13-20

8-10
8-13
8-17
8-10
7-17
10-13
7-13
7-10
7-10
7-10
7- 8
7-13
7-13

X 17-23
X 17-26
X 20-23
X 20-26
X 13-26
X 17-20
17-26
17-23
17-23
X 17-23
X 13-23
X 17-20
X 17-20

X
X
X

Collection No.

T691
T670
T672
T673
T676
T677
T678
T680
T681
T683
T685
T687
T766

Host

B. intermedia
B. intermedia
B. intermedia
B. intermedia
B. interm,edia
B. inlerynedia
B. intermedia
B. intermedia
B. intermedia
B. intermedia
B. intermedia
B. intermedia
B. interm,edia

60

The University Science Bulletin

Ascus-size

Stalk cell -size

in microns

in microns

Collection No. '

Host

40-60 X 13-20

8-10 X 17-20

T690

B.pubescens

33-53 X 13-20

7-17 X 13-20

T692

B. pubescens

46-56 X 15-26

10-23 X 17-30

T643

B. pubescens

50-60 X 15-26

10-17 X 17-26

T635

B. pubescens

43-56 X 13-15

8-10 X 17-20

T620

B. pubescens

33-40 X 15-23

8-10 X 20-27

T619

B. pubescens

33-50 X 12-17

10-13 X 17-23

T618

B. pubescens

33-46 X 13-20

8-10 X 17-23

T480

B. pubescens

36-50 X 12-18

8-17 X 13-23

T335

B. pubescens

T. betulina,

forming typical

witches' brooms.

Ascus-size

Stalk cell -size

in microns

in microns

Collection No.

Host

40-53 X 13-20

8-17 X 17-23

T566

B. aurata

40-63 X 13-23

8-13 X 17-23

T567

B. carpatica

30-53 X 13-20

8-13 X 20-26

T668

B. intermedia

30-50 X 13-20

8-13 X 17-23

T671

B. intermedia

34-46 X 15-21

6-15 X 15-23

T80

B. pubescens

43-63 X 10-17

8-23 X 13-20

T565

B. pubescens

33-50 X 10-20

8-13 X 13-23

T649

B. pubescens

43-56 X 17-26

13-23 X 10-26

T656

B. pubescens

36-46 X 13-23

10-27 X 13-27

T168

B.pubescens

This would give as dimensions for T. lapponica: Asci, 23-73[ji, X
12-23IJ.; stalk cells, l-33\i X 13-27iji,; for T. betulina: Asci, 30-63pL X
10-26[;,; stalk cells, 7-27[x X 10-27[j..

The writer's collections of T. lapponica on B. intermedia and B.
pubescens were made from trees bearing typical witches' brooms
of T. betulina as well as from adjacent trees showing only "shoot-
infections." This was in the exact locality in which Jiiel had col-
lected his fungus. It was apparent that the lapponica-'miections,
which included cases where one or a few leaves of a shoot were
diseased, were juvenile stages of witches' brooms. A witches' broom
being a perennial structure, and only identifiable as a witches'
broom after adventitious buds have developed twigs, it could only
become established through such juvenile stages. One or more
leaves must become infected, mycelium invade the whole shoot, and
the formation and development of adventitious buds be induced.
In localities such as Lappland, where infestation is great and where
infection seems to occur every year, juvenile stages are abundant.

Complete proof of this idea of the formation of a witches' broom
could be furnished by inoculation-experiments, which have not yet
been possible.

c. Taphrina lagerheimii Palm

This fungus was described by Palm as affecting its host {B.
pubescens) in the same manner as T. lapponica but having asci,
53-60[jL X 13-5-17t;.; and stalk cells, 19.5-23.5[x X 19.5-5-26[jl.

Mix: Monograph of Genus Taphrina 61

Palm preserved no type specimens of his various new species of
Tayhrina. The specimens (listed above) studied by the writer
were found by Dr. Th. Arwidsson among Lagerheim's collections
in Stockholm's Hogskola. They had been collected by Palm and
evidently studied by him but bore no date. No collections from
Kapellskar in Uppland was found, that being Palm's type locality
for T. lagerheimii.

Study of these specimens showed dimensions to be: Asci, 33-60[ji
X 10-18[;,; stalk cells, 10-17[jl X 23pL. The stalk cells seen were
shorter and narrower than those reported by Palm but even his di-
mensions fall within the limits for T. betulina (including T. lap-
ponica and T. turgida) : Asci, 23-73[x X 10-26[jl; stalk cells, 7-27\i. X
10-27pL. Clearly T. lagerheimii can not be distinguished from T.
lapponica and it must become synonymous with T. betulina.

d. Taphrina willeana Svendsen

This fungus, described by Svendsen as affecting leaves (wholly or
in part) but not discoloring or thickening them, and not forming
witches' brooms, is obviously like T. lapponica. Svendsen reported
its dimensions as: Asci, 40-50pi, X 15-20^1.; stalk cells, 20-24[jl X
24-30[ji.. Dimensions obtained by the writer from a study of Svend-
sen's type-specimen are: Asci, 33-56t;, X 13-17[x; stalk cells, 10-24[ji,
X 13-24[JL. These dimensions do not distinguish Taphrina willeana
from T. betulina nor from Juel's T. lapponica, nor has it any other
distinguishing feature. It must be included in Taphrina betulina.

Lind (1913) reports the collection by Rostrup of Taphrina betu-
lina on Betula pubescens Ehrh. var. urticifolia (Loud.) Schelle, and
on B. nigra L. These specimens have not been seen by the writer.

36. Taphrina splendens Palm
Taphrina splendens Palm, Arkiv. Bot. 15:1-41. 1917.

Affecting leaves of Betula pubescens Ehrh. Leaves (in specimens
seen) deeper green than normal above, rusty tan beneath, not thick-
ened or enlarged. Apparently not forming witches' brooms.

Mycelium subcuticular.

Asci hypophyllous, cylindric, rounded at the apex; stalk cell
rounded or truncate below, of nearly the same width as the ascus.
Ascospores, eight, ovate to elliptic, frequently budding in the ascus.
(Fig. 14, F, Fig. 15, C.)

Dimensions: Of asci, 50-96[i, X 13-23[a; of stalk cells, 10-33^;- X
10-26[ji.; of spores, 4-6.5[jl X 3-5. Spi,.

62 The University Science Bulletin

Distribution: Northern Norway, Swedish Lappland.

Material examined: Norway: Nordland, Bjelladalen in Nord
Rana, Juy 21, 1926, I. J0rstad. Sweden: Lappland, Abisko, B.
Pahn (presumably the type, though not so designated) ; Lulea, July
12, 1926, J. Vleugel.

Some doubt is felt whether to regard this fungus as a separate
species or to consider it a northern variant of Taphrina betulina. It
is possible that it may, on further study, be found to cause witches'
brooms. Its asci are definitely larger than those of T. betulina.

The longest asci observed in Palm's specimen measured SOpL.
Longer asci (96ijl) were found in J0rstad's collection. If Taphrina
splendens is to be recognized as a separate species it should include
all northern forms with large asci.

37. Taphrina nana Johanson

Taphrina nana Johanson, Ofvers. of K. Svensk. Vetensk. Akad. Forhandl.

1885 :29-47. 1886.

J^xoascus mmus (Johans.) Sadebeck, Jahrb. Hamburg. Wissensch. Anst.

10:5-110. 1893.

Taphrina alpina Johanson, Bih. K. Svensk. Vetensk. Akad. Handl. 13:3-28.

1887.

Exoascus alpinus (Johans.) Sadebeck, Jahrb. Hamburg. Wissensch. Anst.

10:5-110. 1893.

Causing yellowing of leaves, without enlargement or thickening,

affecting part or all of a shoot, or inducing definite witches' brooms

on Betula ermani Cham., B. intermedia Thom., B. japonica Sieb.,

B. nana L., B. pendula Roth., B. pubescens Ehrh.
Mycelium intercellular (subcuticular in early stages).

Asci at first hypophyllous, later amphigenous, cylindric, rounded
or truncate at the apex, with a broad stalk cell seated on the epi-
dermis or v/ith a narrower, wedge-shaped stalk cell inserted be-
tween epidermal cells. Ascospores eight, round, ovate or elliptic,
often budding in the ascus. (Fig. 16, A-C.)

Dimensions: Of asci, 13-30[x X 8-15[jt,; of stalk cells, 7-16[x X
8-20[jl; of ascospores, 3.5-6pi. X 3.5-5ij,.

Distribution: Sweden, Germany, Russia, Kamchatka.

Material examined: B. nana. Norway: S0r Tr0ndelag, Knut-
sh0 in Opdal, July, 1887, A. Blytt. Sweden: Jamtland, Areskutan,
July 12, 1884, C. J. Johanson (F. Par. Scand. Exs. 231b. Labelled
"T. carnea" but apparently duplicate of type) ; ibid., July 16, 1885,

C. J. Johanson (Herb. Bot. Mus. Stockholm 3649) ; between Hallen
and Bydalen, Aug. 3, 1909, 0. Juel; Harjedalen, Glan, G. Lager-

Mix: Monograph of Genus Taphrina

63

Fig. 16. A-C, asci of Taphrina nana. A, on Be tula imna; B, on B.
nana (as T. alpina) ; C, on B. pubescens. B, asci of Taphrina boycei on
Betula jontinalis. All X 900.

64 The University Science Bulletin

heim; Lappland, Abisko, July 28, 1911, 0. Juel (three collections,
two labelled *T. alpina" the other "T. nana var. hyperborea" Juel) ;
ibid., B. Palm; ibid., Aug. 18-21, 1939, A. J. M. (four collections,
three identifiable as T. alpina, one as T. nana).

B. pendula. Germany: Westphalia, vicinity of Siegen, May-
June, 1920, A. Ludwig (Myc. Germ. 1649, as T. turgida). Poland:
Anin, July, 1938, H. Juraskowna (Herb. Inst. Phytopath. Schol.
Sup. Agric. Varsaviensis, as T. turgida). Sweden: Falun, July 6,
1904 (as T. alpina).

B. pubescens. Sweden: Lappland, Abisko, B. Palm.

The distinction between Taphrina nana and T. alpdna, made by
Johanson (1887) and concurred in by Juel (1909) was that T. nana
forms intercellular mycelium throughout the leaf parenchyma, while
T. alpina has subcuticular mycelium. The occurrence of inter-
mediate forms makes this distinction untenable. Specimens identi-
fied by Johanson and by Juel as T. alpina show subcuticular
mycelium, but modification of one or two rows of the spongy paren-
chyma as well as of the epidermis occurs. The asci in these speci-
mens are hypophyllous. A specimen collected by Juel in Jamtland,
Aug. 3, 1909, and identified by him as T. nana, and a specimen
collected at Abisko by the writer Aug. 19, 1939, show modification
of the lower half of the spongy parenchyma, a little intercellular
mycelium, and hypophyllous asci intermediate in size (asci 17-26[x
X 8-10[jL, stalk cells 7-lOp, X 8-13) between those of T. alpina and
T. nana (Johanson's measurements). In most specimens of T. nana
the mycelium extends throughout the leaf, all the leaf tissues are
modified, and the asci are amphigenous, being slightly larger when
occurring on the upper surface of the leaf. The asci of Taphrina
alpina are described by Johanson as 20-27[jl X 9-14[jl, with stalk
cells 8-14 (18)pi. X 12-20[x; the asci of T. nana being 18-30tx X
7-9[x, stalk cells 7-15pi. X 7-17pi.. Actually there is no size-distinc-
tion between the two forms, though the hypophyllous asci (of either
form) are somewhat smaller than the epiphyllous asci.

It is evident that Taphrina alpina is merely a juvenile stage of
Taphrina nana. Infection apparently occurs through the lower epi-
dermis, the spongy parenchyma is first invaded and larger asci form
on the upper epidermis. Since the two species are indubitably syn-
onymous the name Taphrina nana Johanson has priority. It is in-
teresting that in Taphrina nana as in T. betulina early stages can
be found in which one or all of the leaves of a shoot are attacked
but no witches' brooms have yet been formed.

Mix: Monograph of Genus Taphrina 65

The occurrence in Kamchatka of T. alpina on Betula ermani, and
of T. nana on B. japonica are reported by Jaczewski (1926). Speci-
mens were not obtainable.

38. Taphrina americana Mix
Taphrina americana Mix, Trans. Kansas Acad. Sci. 50:77-83. 1947.

Causing witches' brooms on Betula fontinalis Sarg., B. lutea
Michx., and B. papyrifera Marsh.

Mycelium subcuticular, perennial.

Asci hypophyllous, cylindric, rounded or truncate at the apex;
stalk cell broader than the ascus, short and rounded, truncate at
the base; ascospores eight, often budding in the ascus. (Fig. 13,
G-J.)

Dimensions: Of asci, 17-40ijl X 8-18tx; of stalk cells, 6-20[;. X
7-23[jl; of spores, 4-6pL X 3.5-5. 5ix.

Distribution: North America (occasional).

Material examined: B. fontinalis. Colorado: Larimer Co., tribu-
taries South Fork Cache Le Poudre River, Happy Hollow, July 2,
1896, L. H. Pammel (Herb. Iowa Agric. Coll. Plants N. Colo. 118) ;
Boulder Canyon, El Vado, July 27, 1942, A. J. M. (Type).

B. lutea. New Hampshire: Mt. Washington, Alpine Garden, R.
Thaxter (Type).

B. papyrifera. New Hampshire: Mt. Washington, Alpine Gar-
den, July, 1886, R. Thaxter (Type). Wisconsin: Solon Springs,
July 20, 1914, J. J. Davis.

This fungus is considered distinct from both Taphrina betulina
and T. nana because it is intermediate in ascus-size between these
two fungi. It further differs from them in host range and geo-
grapic distribution.

39. Taphrina boycei Mix
Taphrina boycei Mix, Amer. Jour. Bot. 26:44-48. 1939.

Causing small unthickened spots, pale yellow on both sides (re-
sembling those caused by T. flava) on leaves of Betula fontinalis
Sarg., and B. occidentalis *" Hook.

Mycelium intercellular.

Asci hypophyllous, oblong to broad-cylindric, rounded at the apex,
often broadened at the base, arising directly from intercellular my-
celium, containing persistent pale yellow oil globules; ascospores
eight, round, ovate or elliptic. (Fig. 16, D.)

6. Rehder assigns this species in part to Betula fontinalis Sarg. and in part to B. papy-
rifera Marsh, var. occidentalis (Hook) Sarg. Perhapa the form in question belongs to the
latter.

5—3659

66 The University Science Bulletin

Dimensions: Of asci, on B. jontinalis, 30-60[x X 20-36ij,. en B.
occidentalis, 30-46pL X 20-33pL; of ascospores, 4-5[;, X 3.5-4[i.

Distribution: British Columbia.

Material examined: B. jontinalis. British Columbia: near
Cheekye, July 19, 1931, J. S. Boyce and J. L. Mielke (Herb. J. S.
B. 1938. Type).

B. occidentalis. British Columbia: New Westminster, Green
Timbers Forest Nurseiy, July, 1938, Malcolm Wilson; Revelstoke,
Big Bend Highway, June 30, 1931, J. S. Boyce and J. L. Mielke
(Herb. J. S. B. 1963. Type).

40. Taphrina carnea Johanson

Taphrina carnea Johanson, Ofvers. of K. Svensk. Vetensk. Akad. Forhandl.

1885 : 29-47. 1886.

Exoas^cus carneus (Johans.) Sadebeck, Jahrb. Hamburg. Wissensch. Anst.

10:5-110. 1893.

Taphrina janus (Thomas) Giesenhagen, Bot. Zeit. 59:115-142. 1901.

Exoascus janus Tliomas, Forstl. Naturw. Zeitschr. 6:305-314. 1897.

Taphrina lata Palm, Arkiv. Bot. 15:1-41. 1917.

Causing thickened, yellowish to red, leaf-curl lesions on leaves
of Betula fruticosa Pall., B. glandulosa Michx., B. humilis Schrank.,
B. intermedia Thom., B. lutea Michx., B. nana L., B. papyrifera
Marsh., B. pendula Roth., B. pubescens Ehrh.

Mycelium intercellular.

Asci epiphyllous, hypophyllous or amphigenous, broad-cylindric
or oblong, sometimes irregular in outline, rounded or truncate at
the apex, frequently broadened toward the base, sometimes com-
pressed at the base by adjacent asci, lacking a stalk cell. Asco-
spores rarely seen, asci commonly filled with blastospores which
may be small and ovate to elliptic, or minute and bacterioidal.
(Fig. 17, A-C, F.)

Dimensions: Of asci, 30-86;;. X 10-26ij. ; widening at base to as
much as 30i;,; of ascospores, 5-6[a X 3-4. S;;.; of blastospores, 3-6[x X
2-4[jL, or if bacterioidal, 1-6ijl X l-1.5i;..

Distribution: Switzerland, Sweden, Russia, eastern North Amer-
ica.

Material examined: B. glandulosa. New Hampshire: Mt.
Washington, Lake of the Clouds, July 3, 1886. [Identified by Pat-
terson (1985) as T. bacteriosperma.] New York: Mt. Marcy, July,
1895, C. H. Peck. [Identified by Peck (1896) as T. bacterio-
sperma.]

B. humilis. Poland: Janow near Lwow, May 30, 1910, M. Raci-
borski (Myc. Polon. 51).

Mix: Monograph of Genus Taphrina

67

Fig. 17. Asci (X900) of: A, Taphrina carnea on Betula lutea; B,
B. pubescens; C, B. nana; D, T. bacteriosperma on B. pubescens (as
T. janus); E, B. pubescens; F, T. carnea on B. glandulosa; G, T. bac-
teriosperma on B. nana.

68 The University Science Bulletin

B. intermedia. Norway: Finnmark, Skadevarre in Alta, July
30, 1924, I. J0rstad; Troms, Alterdet in Kvaenangen, July, 1900,

C. J. Svendsen. Sweden: Harjedalen, Fjallnas, June 23, 1933, A.
G. Eliasson; Jamtland, Areskutan, July 20, 1892, Juel; Jamtland,

Foot of Snashogarna, July 20, 1889, E. H g; Storlien, Aug.

24, 1939, A. J. M.; Lappland, Vassijaure, Aug., 1903, T. Vestergren
(Micr. Rar. Sel. 721b) ; Palnoviken, Aug. 20, 1939, A. J. M.

B. lutea. Maine: Aquossoc, July 24, 1935, H. B. Peirson (U. S.

D. A. Div. For. Path. 35289). Ontario: Lake Temagami, Meta-
gama Point, June 27, 1929, H. S. Jackson [Univ. Toronto Crypt.
Herb. 1045. Identified by Ray (1940) as T. bacteriosperma]. Que-
bec: Duchesnay, June 12, 1941, R. Pomerleau; Pontneuf, June 21,
1940, id.

B. nana. Norway: Opland, Storhang in S0r-Fron, July, 1893,
A. Blytt. Sweden: Jamtland, Arestukan, July 20, 1884, C. J. Jo-
hanson (duplicate of type) ; Lappland, vicinity of Peskehaure, July
14, 1938, T. Arwidsson.

B. papyrifera. New Hampshire: Chocorua, May 21, 1911, R.
Thaxter.

B. papyrifera var. minor. New Hampshire: Mt. Washington,
July 3, 1939, D. H. Linder.

B. pendula. Sweden: Uppland, O. Ryd, Aug. 8, 1910, T. Vester-
gren (as T. janus).

B. pubescens. Sweden: Sodermanland, road between Strangniis
and Lund, July 13, 1912, T. Holmgren and T. Vestergren (M. Rar.
Sel. 1618, as T. janus) ; Tungelsta, B. Palm (as T. lata) ; Stockholm,
Uggleviken, June 12, 1895, 0. Juel (labelled "Exoascus bacterio-
spermus Joh — janus Thomas! teste Sadebeck").

Betula sp. New Hampshire: Mt. Washington, Lake of the
Clouds, July, 1897, R. Thaxter; ibid, July 3, 1939, D. H. Linder,
det. D. H. Linder. Quebec: Port Harrison, Aug. 6, 1936, N. Po-
lunin, det. D. H. Linder.

Synonymy: a. Taphrina janus

Thomas (1897) although suggesting that T. bacteriosperma, T.
carnea, and T. janus might actually be forms of the same fungus,
described T. janus as a separate species, distinguishing it from T.
carnea only by its amphigenous asci.

Actually it is not possible to distinguish T. carnea from T. janus.
They are identical in their effect on the host, in size, shape, and
basal compression of asci, and in producing asci upon one or both
surfaces of the leaf.

Mix: Monograph of Genus Taphrina 69

b. Taphrina lata Palm

This fungus was described by Palm (1917) as causing enlarge-
ment and moderate thickening of leaves of B. piibescens, and affect-
ing only young seedlings a foot or less tall. It was found only at
Tungelsta, Sweden, near the railway station. He reported it as
having asci 40-45[jl X 18-22[i,, with a stalk cell measuring 16-20[jl X
25-33[x. The material studied by the writer consisted of a large
packet among Lagerheim's collections, labelled only "Tungelsta,
Palm." It contained a number of small seedling birches, with leaf-
lesions corresponding closely to Palm's description. It seems highly
probable that this is the material Palm used in describing Taphrina
lata.

This is, however, typical material of T. carnea. The asci (lacking
stalk cells) measure 36-60ix X 13-23iJi., widening at the base and
showing compression, exactly as in T. carnea. If this is Palm's
fungus it is difficult to understand how he could have described it
as having a stalk cell. In sections made by the writer asci with
bases inserted and thus overlain by epidermal cells suggested asci
with stalk cells. The mycelium is intercellular, not subcuticular as
stated by Palm.

The occurrence of T. carnea on B. fruticosa Pall. {B. gmelini
Bge.) is reported by Jaczewski (1926).

41. Taphrina hacteriosperma Johanson

Taphrina hacteriosperma Johanson, Bih. t. K. Svensk. Vetensk. Akad.
Handl. 13:3-28. 1897.

Exoascus bacteriospermus (Johans.) Sadebeck, Jahrb. Hamburg. Wis-
sensch. Anst. 10:5-110. 1893.

Causing yellowing of leaves, without thickening but sometimes
with lateral enlargement, often affecting all the leaves of a shoot,
but not causing witches' brooms, on Betula glandulosa Michx., B.
intermedia Thom., B. nana L., B. pubescens Ehrh.

Mycelium subcuticular.

Asci epiphyllous, sometimes also hypophyllous, broad-cylindric,
sometimes irregular, rounded or truncate at the apex, broadened at
the base and often somewhat compressed by adjacent asci, lacking
a stalk cell. Ascospores rarely seen, asci filled with minute bac-
terioidal blastospores (Fig. 17,A-E, G).

Dimensions: Of asci, 33-80[jl X 14-20[x, widening at the base to
23-30[ji.; of ascospores, 3.6-4.5[ji. diam. (Johanson); of blastospores,
3-6[ji, X l-2iJi., or bacterioidal, 1-1. Spi. X O-Six.

Distribution: Norway, Sweden, Greenland, Russia, North Amer-
ica.

70 The University Science Bulletin

Material examined: B. glandulosa. New Hampshire: Mt. Wash-
ington, near Lake of the Clouds, July 3, 1939, D. H. Linder, det.
D. H. Linder.

B. nana. Norway: Dovrefjeld, foot of Mt. Nordra Kunestio,
July 9, 1933, A. G. Eliasson. Sweden: Hiirjedalen, Fjallnas, July
27, 1933, 0. Ostergren; Jamtland, Arestukan, Ulladalen, July 27,
1895, C. J. Johanson (Type).

B. pubescens. Sweden: Lappland, Abisko, July 28, 1911, 0. Juel.

This species is not to be distinguished by the size of its asci from
Taphrina carnea. A distinctive feature is the compression of the
widened basal part of the ascus, where asci stand close together.
Juel (1909) says that the asci are "grown together a little at the
base."

Taphrina bacteriosperma causes no thickening of the leaves of
its host, but often some lateral enlargement. Modification of leaf
tissues is confined to the epidermis and palisade parenchyma; while
T. carnea causes typical thickened "leaf-curl," with all mesophyll
tissues modified. Since such variation in method of attack is pos-
sible within a single species (as indicated earlier for Taphrina nana)
it would be interesting to compare these two fungi further by
means of cultural studies and inoculations. Obviously they are
closely related.

42. Taphrina betulae (Fkl.) Johanson

Taphrina betulae (Fkl.) Johanson, Ofvers. K. Svensk. Vetensk. Akad.
Forhandl. 1885 :29-47. 1886.

Exoasciis betulae Fuckel, Jahrb. Nassau. Verein. Naturk. 27 and 29:1-99.
1873 and 1874.

Ascomyces betulae, Magnus, Rab. F. E. 2734.
Taphrina auctumnalis Palm, Arkiv. Bot. 15:1-41. 1917.
Taphrina betulae (Fkl.) Johans. var. auctumnalis Sadebeck, Jahrb. Ham-
burg. Anst. 10:5-110. 1893.

Causing small (up to 1 cm. diam.) pale green to yellow (brown
in age) unthickened spots on leaves of Betula intermedia Thom.,
B. medwediewi Reg., B. pendula Roth., and var. purpurea Schneid.,
B. pubescens Ehrh., and B. turkestanica Litvin. Spots are evident
on either surface of the leaf, showing as a pale area on the op-
posite surface.

Mycelium, subcuticular.

Asci epiphyllous, hypophyllous or amphigenous, cylindric, rounded
or truncate at the apex, sometimes broadened at the base. Stalk
cells seated, broad and flat, broader than the ascus, or nearly iso-
diametric, not broader than the ascus. Ascospores eight, ovate to
elliptic, sometimes budding in the ascus. (Fig. 18, A-E.)

Mix: Monograph of Genus Taphrina

71

Fig. 18. Asci (X900) of A-E, Taphrina betulae. A, on Betula pen-
dula; B, B. pubescens; C, B. intermedia; D, B. pubescens (as T. auc-
tumnalis) ; E, B. medwediewi; F, T. flava on B. papyrijera.

72 The University Science Bulletin

Dimensions: Of asci, 17-46[ji, X S-lSt*- ; of stalk cells, 7-17[x X
8-30pi,; of ascospores, 4-6[jl X 3.5-5n.

Distribution: northern Europe.

Material examined: B. intermedia. Sweden: Lappland, Aug.
21, 1939, A. J. M.

B. medwediewi. Russia: Causasus, Mt. Anczcha, Aug. 16, 1917,
W. Siemaszko (Herb. W. S.).

B. pendida. Germany: Berlin, Thiergarten, June 12, 1880, P.
Magnus (Rab.-Wint. F. E. 2734). Sweden: Grisslehamn, Sept. 9,
1912.

B. pendula var. purpurea. Germany: Berlin, Spath'sche Baum-
schule, Sept., 1891, P. Sydow (Myc. March. 3358).

B. pubescens. Germany: Berlin, Lichterfelde, Oct., 1895 (M.
March. 4337, as var. auctumnalis) ; Hamburg, Wandsbek, Sept.,
1891, Sadebeck (Munich Herb. Type of var. auctumnalis) ; O.
Schlesien, Werdenau, June, 1913, J. Horuby (Petrak. Fl. Boh. Mor.
Exs. Pilze 776). Norway: Bjsokeny, Aug., 1893, G. Lagerheim (as
var. auctumnalis).

B. turkestanica. Germany: Berlin, Spath'sche Baumschule,
Sept., 1898, P. Sydow (Myc. March. 3359).

Betula sp. Sweden: Sodermanland, Tungelsta, B. Palm.

Sadebeck (1893) described T. betulae var. auctumnalis on the
basis of somewhat smaller asci, 15-27[Jt. X 6-9[ji,; stalk cells 2-5pi. high
and very broad, stating that it occurred on B. pubescens and rarely
on B. pendida. Palm in raising this variety to specific rank, con-
fined it entirely to B. pubescens, calling the form on B. pendula
T. betulae. There seems to be no basis for separation of the fungus
here studied from T. betulae either as a variety or as a separate
species, dimensions of asci and stalk cells falling well within those
observed for T. betulae. Asci as short as 15[x or stalk cells as short
as 5tJL were not seen in Sadebeck's type material. Presumably, since
this is an autumn form, it results from late primary or from sec-
ondary infection.

43. Taphrina jiava Farlow

Taphrina flava Farlow, Proc. Amer. Acad. Arts and Sci. 18:65-85. 1883.
Magnusiella flava (Farl.) Sadebeck, Jahrb. Hamburg. Wissensch. Anst.
10:5-110. 1893.

Causing small (up to 5 mm.) yellow to brown or red-brown spots
on leaves of Betula papyrifera Marsh., and B. populifolia Ait.
Mycelium intercellular.

Mix: Monograph of Genus Taphrina 73

Asci hypophyllous at first, later amphigenous, oblong (on upper
epidermis) to squarish (on lower epidermis), arising from the ends
of narrow hyphae. Ascospores rarely seen, asci filled with ovate to
elliptic blastospores. (Fig. 18, F.)

Dimensions: Of asci, 30-63[j. X 15-33ix; of blastospores, 5-6[x X
5-5.5[JL, or minute.

Distribution: eastern North America.

Material examined: B. papyrifera. New Hampshire: Mt.
Washington, carriage road, E. Faxon.

B. populijolia. Maine: Kittery Point, July 7, 1929, J. H. Faull
(Herb. J. H. F. 9223). Massachusetts: Newton, W. G. Farlow
(N. A. F. 300) ; Roxbury, July 4, 1888, A. B. Seymour (E. F. 171) ;
Waverly, June 17, 1902, A. B. Seymour. New Hampshire: Mt.
Washington, head of Great Gulf, July, 1886, W. G. Farlow. Nova
Scotia: Lunenberg Co., Chester, July 22, 1929, J. H. Faull (Herb.
J. H. F. 9252).

44. Taphrina robinsoniana Giesenhagen

Taphrina robinsoniana Giesenhagen, Flora 81:267-361. 1895.

ExoasciLS robinsonianus Saccardo and Trotter, Sylloge Fungorum 22:765.

Taphrina rugosa Ray, Mycologia 31:56-75. 1939.

Causing protruding tongue-like enlargements of bracts and ova-
ries of female catkins of Alnus incana Moench., and A. rugosa
Spreng. The tongues appear on the catkins near pollination time
and are then very small: 2-3 mm. long. Later in the season when
the fruits of the alder are well grown the tongues may measure sev-
eral millimeters in length. Occasionally also causing leaf curl
(involving all or part of the leaf), with typical thickening of leaves,
on the same hosts.

Mycelium intercellular.

Asci occurring all over the surface of the tongue, amphigenous
on leaves, cylindric, rounded or truncate at the apex, with a stalk
cell which may be broader than the ascus and rather long. Asco-
spores eight, ovate to elliptic, often budding in the ascus, producing
numerous ovate to elliptic blastospores. (Fig. 19,A-D.)

Dimensions: Of asci, 13-43[jl X 6-17[jl; of stalk cells, 6-20\l X
5-17[ji.; of ascospores, 2.5-6[ji. X 2.5-5[jl.

Distribution: eastern North America.

Material examined: A. incana. Connecticut: Collinsville, Aug.,
1926, A. J. M.; East Granby, Dec. 13, 1908, P. Spaulding (U.S.
D. A. Div. For. Path. 2028). Maine: Byron, Aug. 25, 1938, A. E.

74

The University Science Bulletin

Fig. 19. Asci (X900) of A, Taphrina robinsoniana on Almis incana; B,
C, A. rugosa; D, the same (T. rugosa); E, T. sadebeckii.

Mix: Monograph of Genus Taphrina 75

Prince; Ellsworth, Sept. 14, 1937, J. R. Hansbrough (U. S.D.A.
Div. For. Path. 82125) ; Eustis, July 19, 1935, J. R. Hansbrough
(Herb. J. R. H. 2385) ; Mt. Desert, Bar Harbor, J. S. Boyce (Herb.
J. S. B. 2026); Hamden, July 5, 1938, A. E. Prince. Massa-
chusetts: Amherst, Aug., 1934, R. H. Thompson; Granville, Aug.
3, 1885, A. B. Seymour (Econ. F. 167a) ; Jamaica Plain, Arnold
Arboretum, July 28, 1892, A. B. Seymour. Michigan: Copper
Harbor, Aug., 1928, A. J. M.; Empire, Aug. 10, 1938, E. A. Bessey.
New Hampshire: Carroll, Cherry Mountain, July 27, 1926, P.
Spaulding (For. Path. 45312). New York: Blue Mt. Lake, Aug.
24, 1934, G. E. Thompson (Cornell Univ. Dept. Plant Path. Herb.
23806) ; Ithaca, Ellis Hollow, June 22, 1940, A. J. M.; ibid., July 15,
1940,id.; ibid., July 23, 1940, id. North Carolina: Old Fort, Oct.
6, 1930, P. Spaulding. Vermont: Bethel, Sept., 1913, P. Spaulding
(For. Path. 16833). New Brunswick: Fredericton, July 18, 1936,
I. L. Connors (Cent. Expt. Farms Herb. 4518). Newfoundland:
Harpoon River, Aug. 18, 1928, P. Spaulding (For. Path. 45640) ;
Spruce Branch, Aug. 23, 1928, P. Spaulding (For. Path. 45649).
Nova Scotia: Mahione Bay, Covey Island, July 20, 1936, H. T.
Giissow (Cent. Expt. Farms Herb. 4527). Ontario: Lake Tema-
gami. Bear Island, Aug. 10, 1930, I. L. Connors (Cent. Expt. Farms
Herb. 1607) ; Sault Ste. Marie, Sept. 10, 1940, A. J. M. Quebec:
Deschambault, I. L. Connors (Cent. Expt. Farms Herb. 1137) ;
Farnham, Aug. 11, 1934, H. N. Racicot (Ottawa Div. Bot. Herb.
3727) ; Gaspe, Aug. 7, 1894, F. F. Forbes (Phanerogam. Herb. Stan-
ford Univ.).

Alnus rugosa. Alabama: Auburn, April, 1891, G. F. Atkinson
(Econ. F. 167b). Arkansas: New Hope, Oct. 15, 1932, D. Demaree
(Phanerogam. Herb. Stanford Univ.). Connecticut: Collinsville,
Aug. 23, 1937, A. J. M. Georgia: Athens, July 8, 1940; G. E.
Thompson; Crawford, Oct. 17, 1935, J. H. Miller (Univ. of Georgia
Herb. 2772) ; Experiment, April 9, 1937, W. A. Jenkins (Cornell
Univ. Dept. Plant Path. Herb. 27346. Ray's type of Taphrina
rugosa); ibid., April 2, 1940, W. A. Jenkins; ibid., April 8, 1940,
id.; ibid., April 22, 1940, id.; ibid., May 2, 1940, id.; ibid., May 20,
1940, id.; ibid., Aug. 14, 1940, id. Massachusetts: Quincy, July,
1939, D. H. Binder (on both catkins and leaves) ; ibid., Aug. 9, 1940,
id. New York: Ithaca, June 23, 1940, A. J. M.; ibid., July 12, 1940,
id.; ibid., Aug. 9, 1940, id. Tennessee: Montvale Springs, April 15,
1934, A. J. Sharp and L. R. Hesler (Univ. Tenn. Herb. 6094). Vir-
ginia: Fairfax Co., Black Pond, July 12, 1922, A. E. Jenkins.

76 The University Science Bulletin

Alnus sp. Massachusetts: Manchester, July 17, 1916, W. G.
Farlow; Stony Brook, Oct., 1893, E. A. Burt; Wellesley, July 23,
1935, D. H. Linder; Worcester, July, 1924, H. E. Greenwood.

Ray (1939) described as a new species Taphrina rugosa on very
small catkins of Alnus rugosa, using material collected by W. A.
Jenkins at Experiment, Georgia. The distinguishing features were
the very small, slender tongues, 2 or 3 mm. long, and the small asci
(asci, 14-28pL X 4.5-9[x, stalk cells, 8-14[x X 6-lOix) and the small
spores (2-4.5[x diam.) which were often 4 or 6 rather than 8 per
ascus. Ray stated this to be a southern form, distinct from the
common Taphrina robinsoniana of the northeast.

Since it seemed possible that Taphrina rugosa might be a juvenile
stage of T. robinsoniana, probably to be found on both Alnus incana
and A. rugosa, special study of the question was undertaken. In
examination of Ray's type material no asci were seen so short as
those reported by him, the dimensions observed being: Asci, 23-33ijl
X 8-lOpi, ; stalk cells, 10-17[x X 5-8pi,. Shorter asci were, however,
observed later in certain other specimens.

Following examination of the type specimens a search was made
among various collections for specimens of Alnu^ incana or A. rugosa
showing very small tongues. The following were found: A. incana.
Spruce Branch, Newfoundland, Aug. 23, 1928, P. Spaulding, tongues
2-3 mm. long, asci 16-36pi, X 8-10[J., stalk cells 6-17ii. X 8-13[ji,;
Hamden, Maine, July 5, 1938, A. E. Prince, tongues 2-3 mm. long,
asci 17-23[iX8-10[ji,, stalk cells 7-13[xX5-10[ji. A. rugosa, Montvale
Springs, Tennessee, April 15, 1934, A. J. Sharp and L. R. Hesler,
tongues 2-3 mm. long, asci immature, only ascogenous cells being
present; Black Pond, Virginia, July 12, 1922, A. E. Jenkins, tongues
5-8 mm. long, asci 20-30[;, X 7-10ijl, stalk cells 10-17[x X 7-10[x.

During a stay at Cornell University in 1940 the writer received
through the kindness of Dr. W. A. Jenkins specimens of "Taphrina
rugosa" on Alnus rugosa collected by him on April 2, April 22,
May 2, May 20, July 8, and August 14. In the first two of these
collections only a few very short tongues were evident and the
fungus was immature. Doctor Jenkins suggested (in a letter of
April 9) that the overgrowths were from enlarged ovaries rather
than from bracts. This is true of the earliest outgrowths observed
but in the material collected April 22 and later, as well as in the
specimens discussed above, both ovaries and bracts were enlarged.
Contiguous bracts and ovaries may show tongues.

In the collection of April 22 the tongues were 2-3 mm. long, and

Mix: Monograph of Genus Taphrina 77

a few mature asci were present, measuring 26-33[ji. X 7-lOti,, stalk
cells 10-13[x X 8-lOpi,. In the collection of May 2, the tongues were
5-10 mm. long and 2-3 mm. wide. On tongues from ovaries asci
were 13-33pL X 7-10[i, stalk cells 18-17[;, X 7-10[ji,; on those from
bracts asci measured 23-40iJt. X 7-10[jl, stalk cells 13-17pi. X 7-10[jl. In
the collection of May 20 the tongues were 1 to 3 cm. long, although
the catkins were still very small. Asci measured 26-36[ji, X 8-10[j,,
stalk cells 8-20[i. X 8-13ij,.

It is to be noted that in the earliest collections showing mature
asci some of these asci were large enough to be characteristic of
Taphrina robins oniana. Ascospores measured 4.5-5ix X 4-4. SpL and
were frequently 4 or 6 per ascus rather than 8. In the specimens of
May 20, eight-spored asci were more common and some asci con-
tained blastospores.

In the collections of July 8 and August 14 the outgrowths were
long (typical of T. robinsoniana) and asci measured 26-32tJL X 7-9ijl;
stalk cells 8-13[l X l-lS[i.

Alnus rugosa is rare in the vicinity of Ithaca but a few trees
occur in Stewart Park along Cayuga Lake Inlet. These were vis-
ited on May 14, 1940, when the male catkins were shedding pollen.
A week later pollination was over although styles and stigmas were
still evident. There was no sign of fungus attack. Observations
were continued at weekly intervals and on June 23, a number of
reddish tongues, about 2 mm. long, were seen protruding from some
catkins. These catkins were still very small, about "pollination-
size," styles and stigmas were still evident, and the catkins were
quite like those in the early collections received from Georgia.

Weekly visits to these alders were continued and the gradual
enlargement of the outgrowths watched. It became evident that
the large "summer" tongues characteristic of T. robinsoniana are
enlarged states of the very small early outgrowths.

Collections were made on June 23, July 12, and August 9. On
July 12 infected leaves were also collected. These collections may
be summarized as follows:

June 23, tongues 2 mm. long, asci 20-26[i X 8-10[jl, stalk cells 8-10[x
X 7-10[jL, spores often 6 per ascus, 4.5[jl X 4(ji..

July 12, tongues 5 mm. long, asci 26-40[jl X 8-12pi., stalk cells 8-20[ji.
X 8-12[jL, spores often 8 per ascus, 4.5-5[jl X 8-12[jl, blastospores pres-
ent.

August 9, tongues 2 cm. long, asci 26-36[x X 8-12[ji,, stalk cells
10-17[x X 8-12[A, spores mostly blastospores.

78 The University Science Bulletin

On the curled leaves collected July 12, the amphigenous asci
measured 17-26[ji, X 7-10[a, stalk cells 7-lOpi, X 7-10[jl. This is in agree-
ment with observations made on material collected at Quincy, Mas-
sachusetts, July, 1939, by D. H. Linder. In these specimens asci
from catkins measured 23-40i;, X 8-12pL, stalk cells 13-20[;, X 7-13^/,.
asci from leaves 23-26[x X 7-10[i.; stalk cells 10-17[j. X 1-l^v- Evi-
dently asci formed on leaves are regularly somewhat smaller than
those on catkins, a fact already noted for T. robinsoniana by Ray
(Lc).

Coincident with these observations of the fungus on Alnus rugosa,
its development on A. incana was watched. Specimens of these
alders growing at Ellis Hollow, near Ithaca, were inspected at fre-
quent intervals beginning May 15. Collections were made on June
22, July 15, and July 23. Obsei-vations were as follows:

June 22, tongues 1-2 mm. long, asci 20-33[ji, X 8-10[x, stalk cells
8-17tJ. X 8-13[x, spores mostly 8 per ascus, 4-6[x X 4-4.5[ji,.

July 15, tongues 1 cm. long, asci 17-36[j. X 8-10[i-, stalk cells 8-16[x
X 8-13[;., spores 5-6pL X 4.5-5[ji,, mostly blastospores.

July 23, tongues 1-1.5 cm. long, asci 26-43iJt, X 10-13[/., stalk cells,
8-17[ji. X 7-10[i,, spores mostly blastospores.

Infected leaves were collected on July 23. On these the asci were
somewhat smaller than on the catkins.

From these observations two conclusions are inescapable: that
there is no morphological distinction between Taphrina robinsoniana
on Alnits incana and on Alnus rugosa, and that Taphnna rugosa is
a juvenile stage of T. robinsoniana.

American collectors have been slow to recognize the identity of
Taphrina robinsoniana and many collections have been reported as
T. alni-incanae (Kiihn) Magn. [T. amentormn (Sadeb.) Rostr.].
Conners (1932) has mentioned this error in identification and has
suggested the probability that Taphrina amentorum does not occur
in eastern North America. Conners' suggestion led to the exami-
nation of a very large number of collections (listed above). All
specimens proved to be Taphnna robinsoniana. Taphrina amen-
torum on Alnus rubra has been reported from Alaska (Ray, 1939).

45. Taphrina sadebeckii Johanson

45. Taphrina sadebeckii Johanson, Ofv. of K. SVensk. Vetensk. Akad. Forhandl.
1885:29-47. 1886.

Exouscics alni de Bary in Litt., in part.

Exoascus flavus Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 1:93-124.
1884.

Exoascus epiphyllus Sadeb. var. maculans Sadebeck, Jahrb. Hamburg. Wis-
sensch. Anst. 8:61-95. 1890.

Mix: Monograph of Genus Taphrina 79

Ascomyces tosquinetii Westerdorp, Bull. Acad. Roy. Sci. Lett, et Beaux-
arts Belgique 2 ser. 11:644-660. 1861, in part.

Causing moderate-sized (up to 1 cm. diam.) yellow spots on
leaves of Almis glutinosa Gaertn., A. hirsuta Turcz., A. hirsuta
Turcz. var. sibirica Schneid., and A. nigosa Spreng.

Mycelium intercellular.

Asci hypophyllous, sometimes also epiphyllous, cylindric, rounded
or more often truncate at the apex, often with pale yellowish epi-
plasm, inserted in a broad stalk cell (often broader than the ascus).
Ascospores eight, ovate to elliptic, frequently budding in the ascus.
Ascus emerging from the chlamydospore (ascogenous cell) by rup-
ture of the chlamydospore wall. (Fig. 19, E.)

Dimensions: Of asci, 17-65[x X 10-21iJi,; of stalk cells, 7-23[i, X
IS-SOpi.; of ascospores, 4-6tx X 3.5-5[ji,.

Distribution: Europe, Japan.

Material examined: Alnus glutinosa. Denmark: Jylland, Vi-
borg, June 15, 1903, J. Lind. Germany: Brandenburg, Triglitz,
July 20, 1903, 0. Jaap (Myc. Ger. 172) ; Hamburg, Eppendorfer
Moor, July, 1930, A. J. M. Ireland: County Wicklow, Killa-
dreenan House, June 22, 1935, A. J. M. Sweden: Stockholm, Lid-
ingo, July 22, 1884, J. Erikson (F. Par. Exs. 226) ; Rydbo, Aug. 6,
1939, A. J. M.; Statens Vaxtskydds Anstalt, Aug. 12, 1939, A. J. M.

A. hirsuta. Germany: Berlin, Sept., 1891, P. Sydow (Myc.
March. 1360). Japan: Iwate, Mt. Himekami, June 15, 1907, G.
Yamada (Herb. Morioka Imp. Coll. Agric. and For., as T. epi-
phylla) ; Mt. Hayachine, June 13, 1909, id. (as T. epiphylla).

A. rugosa. Sweden: Stockholm, Bergianska Tradgarden, Aug.
10, 1939, A. J. M.; ibid., Aug. 11, id.; ibid., Aug. 12, id.

Sadebeck reports this fungus on A. hybrida A. Br. Three collec-
tions so labelled were found in exsiccati: Krieger, F. Sax. 70 (Kon-
igstein, Aug.-Sept., 1904, W. Krieger), and Herb. K. Stares (Riga,
Latvia, July 30, 1930, Gulbene, Lithuania, Aug. 30, 1931, K. Stares).
In these collections the host seems to be Alnus incana, and the
fungus Taphrina epiphylla. Occurrence of T. sadebeckii in Russia
on A. hirsuta var. sibirica Schneid. is reported by Jaczewski (1926).

Of interest is the occurrence of this fungus in Sweden on the
American alder Alnu^ rugosa. The trees observed were heavily in-
fested, indicating that if Taphrina sadebeckii were introduced into
eastern North America a susceptible host would be waiting.

80 The University Science Bulletin

46. Taphrina tosquinetii (Westend.) Tulasne

Taphrina tosquinetii (Westend.) Tulasne, Ann. Sci. Nat. 5 Ser., Bot. 5:122-
136. 1866.

Ascomyces tosquinetii Westendorp, Bull. Acad. Roy. Sci. Lett, and Beaux-
arts Belgique. 2 Ser. 11 :644-660. 1861.

Taphrina alnitorqua Tulasne, Ann. Sci. Nat. 5 Ser. Bot. 5:122-136. 1866,
in part.

Exoascus alnitorquus Sadebeck, Jahrb. Hamburg Wissensch. Anst. 1 :93-124.
1884, in part.

Exoascus alni de Bary in litt., in part.
Taphrina media Palm Arkiv. Bot. 15:1-41. 1917.

Deforming leaves (leaf-curl of part or whole blade) of Alnus
crispa (Ait.) Pursh. var. mollis Fern., A. glutinosa Gaertn., and A.
hybrida A. Br. Affecting whole shoots or shoot-systems, but not
forming true witches' brooms.

Mycelium subcuticular, perennial.

Asci amphigenous, cylindric, truncate at apex, stalk cells inserted
between epidermal cells. Ascospores eight, often budding in ascus,
ovate to elliptic. Ascus emerging from the chlamydospore (asco-
geous cell) by rupture of the chlamydospore wall. (Fig. 22, B-D.)

Dimensions: Of asci, 17-40pL X 7-13[jl; of stalk cells, 7-17[jl X
8-17[jl; of ascospores, 2.5-5.5[i. X 2.5-5eji..

Distribution: Europe.

Material examined: A. crispa var. mollis. New Hampshire:
Lyme Center, Holt's Ledge, June 26, 1929, H. H. Whetzel and D. S.
Welch (Herb. Cornell Univ. Dept. Plant Path. 17566).

Alnus glutinosa. Belgium: Vallee de Munster (Ht. Rhin.), Aug.
25, 1872 (Herb. Inst. Bot. Strasbourg). Germany: Brandenburg,
between Ruhlsdorf and Marienweder, Aug. 6, 1920, H. and P. Sydow
(Myc. Germ. 1648) ; Hamburg, Eppendorfer Moor, July, 1930,
A. J. M. ; Thuringia, between Jescha and Berka, near Sondershausen,
Sept. 6, 1905, G. Oertel (Myc. Germ. 492). Ireland: County
Wicklow, Killadreenan House, June 23, 1935, A. J. M.; Killarney,
June 19, 1935, A. J. M. Sweden: Gota Canal near TroUhattan,
Aug. 26, 1935, A. J. M.; Radmanso, July 9, 1939, A. J. M.; Tun-
gelsta, July 13, 1939, A. J. M.; Virabruk-i-Rosslagskulle, July 18,
1939, A. J. M. (loc. class, for T. media).

A. hybrida. Germany: Hamburg, Eppendorfer Moor, June 9,
1905, 0. Jaap (F. Sel. Exs. 305b).

Palm (1917) described Taphrina media as intermediate between
T. tosquinetii and T. epiphylla. Apparently he meant to give di-
mensions of asci as 25-30iji. X 10-12ti,, and they are so stated in his
key to species, but in his species-description length of asci is given
as 25-90[ji.. Obviously this is a typographical error; it has been pre-

Mix: Monograph of Genus Taphrina 81

served by later writers. Since Palm left no type specimen of
Taphrina media and no specimen of any fungus collected by Palm
on Alnus glutinosa was to be found among Lagerheim's collections,
a visit was paid on July 18, 1939 to the type-locality, Virabnik.
(This was arranged through the kindness of Dr. T. Arwidsson.)

Virabruk is an abandoned iron foundry, and the alders along the
brook running through it are comparatively few, so that a thorough
search was easily accomplished. The so called "witches' brooms"
pictured by Palm (his figure 4) were easily found, appearing as
upturned clusters of twigs at the ends of branches on the larger
alders. They were entirely free of any fungus, and whatever their
cause, were obviously not due to the action of any species of Taph-
rina. Palm must have collected his fungus from the diseased shoots
of the younger alders nearby. These shoots were attacked by Taph-
rina tosquinetii, and this fungus was the only species of Taphrina
to be found at Virabruk. The collection of T. tosquinetii made
there showed asci 20-30i;. X 8-13pt.; stalk cells, 8-17pL X 10-13[x. The
asci could be considered intermediate between those of T. tosquinetii
and those of T. epiphylla if the dimensions of those fungi as given
in the literature were correct. If the revised dimensions of T. tos-
quinetii (as given above) are considered, the asci clearly belong to
that fungus. It must be concluded that Palm was in error in think-
ing he had a new species and that the fungus he described was
Taphrina tosquinetii.

The fungus on Alnus crispa var. mollis was identified by Dr. A. E.
Jenkins as Taphrina media Palm. A letter from Prof. H. H. Whetzel
to Dr. C. L. Shear is enclosed with the specimens in the Cornell
Herbarium. This states: "The effect on the plant is that of sys-
temic invasion, a sort of witches' broom, the affected leaves standing
up in a staring fashion." This description might apply to the effect
of Taphrina tosquinetii on Alnu^ glutinosa. The asci of the New
Hampshire fungus are shorter than the longest asci observed in T.
tosquinetii, measuring 16-26[x X 8-13[jl; stalk cells 8-13[x X 8-15pL.
The ascus emerges from the chlamydospore in the same manner as
in T. tosquinetii. It seems better to assign this fungus to Taphrina
tosquinetii, even though that fungus is not heretofore known from
North America, than to erect a new species on the basis of this one
collection.

6—3659

82 The University Science Bulletin

47. Taphrina amentorum (Sadeb.) Rostrup

Taphrina amentorum (Sadeb.) Rostrup, Vidensk. Meddel. Naturh. Foren.

Kj0benhavn. 1890:246-264.

Exoascus amentorum Sadebeck, Sitzungsber. Ges. Bot. Hamburg 4:90. 1888.

Aftcomyces alni Berkeley and Broome, Ann. and Mag. Nat. Hist. 17:129-

145. 1876.

Exoascus alnitorquus (Tul.) Sadebeck var. alni-incanae J. Kiihn. (Fungi

Europaei 1616.)

Taphrina alni-incanae (Kiihn) Sadebeck, Jahrb. Hamburg Wissensch. Anst.

8:16-95. 1890.

Exoascus alnitorquus (Tul.) Sadebeck, Jahrb. Hamburg Wissensch. Anst.

1:93-124. 1884, in part.

Exoascus alni de Bary, in litt., in part.

Exoascus alni de Bary var. strobilimis Thiimen, Flora 63:312-322, 323-332.

1880.

Ascomyces tosquinetii Westend. var. strobilina Rostrup, Tidskr. f. Skov-

brug 4:113-206. 1880.

Causing rather large (larger than those caused by T. robinson-
iana) tongue-like outgrowths from the female catkins of Alnus
glutinosa Gaertn., A. hirsuta Turcz., A. hybrida A. Br., A. incana
Moench, and A. rubra Bong.

Mycelium intercellular.

Asci cylindric, truncate at apex, lacking a stalk cell, inserted be-
tween epidermal cells, Ascospores eight, often budding in the ascus,
ovate to elliptic. Asci emerging from the chlamydospore (asco-
genous cell) by rupture of the chlamydospore wall. (Fig. 20, A-C.)

Dimensions: Of asci, 26-53(jl X 10-23ijl; of ascospores, 4.5-6[jl X
4-5[jL.

Distribution: Europe, Japan, Alaska.

Material examined: A. glutinosa. Germany: Mecklenburg,
Warnemlinde, Aug. 16, 1904, 0. Jaap. (F. Sel. Exs. 78a).

A. hirsuta. Japan: Iwate, Gomyojin, Aug. 17, 1907, G. Yamada
(Herb. Morioka Imp. Coll. Agric. and For.).

A. hybrida. Germany: Mecklenburg, Warnemiinde, Aug. 16,
1904, 0. Jaap. (F. Sel. Exs. 78b).

A. incana. Germany: Bavaria, Lechufer near Fussen, Aug. 18,
1912, H. Sydow (Myc. Ger. 1111) ; Fomerania, Ruhgenmahder-
mtinde, Sept., 1894, P. Sydow (Herb. Sydow). Poland: Pieniny,
Aug., 1928, W. Siemaszko (Herb. Inst. Phytopath. Schol. Sup. Ag-
ric. Varsaviensis).

A. rubra. Alaska: Lake Pironsi Glacier, B. E. Fernow (Phan-
erogam Herb. Cornell Univ.).

This fungus has been widely known as Taphrina alni-incanae
(Kiihn) Magnus. As Ray (1939) has pointed out, the first descrip-
tion was by Sadebeck (1888) who called it Exoascus amentorum.

Mix: Monograph of Genus Taphrina

83

Fig. 20. Asci (X900) of A, Taphrina amentoruw. on Alnus glutinosa;
B, A. incana; C, A. hybrida ; D, T. epiphylla on A. incana; E, A. incana
f. aurea; F, G, T. epiphylla {T. klebahni).

84 The University Science Bulletin

Rostrup (1890) transferred it to Taphrina, it thus becoming Taph-
rina amentorum (Sadeb.) Rostrup. Ray (1939) has been the first
to record the occurrence of this fungus on the North American con-
tinent.

48. Taphrina epiphylla Sadebeck

Tnphrina epiphylla Sadebeck, Jahrb. Hamburg Wissensch. Anst. 8:61-95.

1890.

Exoascus epiphyllus Sadebeck, Jahrb. Hamburg Wissensch. Anst. 1 :93-124.

1884.

Taphrina sadebeckii Johans. var. borealis Johanson, Ofvers. K. Svensk

Vetensk. Akad. Forhandl. 1885:29-47. 1886.

Taphrina borealis Johanson, Bih. K. Svensk Vetensk. Akad. Handl. 13:3-28.

1887.

Taphrina klebahni Wieben, Forsch. auf Geb. Pflanzenkr. und Immun. in

Pflanzenr. 3:139-176. 1927.

Causing witches' brooms and (in midsummer) yellowish spots on
leaves of Alnus incana Moench. Leaves not thickened.

Mycelium subcuticular, perennial.

Asci amphigenous, broad-cylindric, usually truncate and often
abruptly widened at the apex to a flat head, inserted in a broad
stalk cell which may be truncate or pointed below. Ascospores eight,
ovate to elliptic, commonly budding in the ascus. Pale yellowish
epiplasm present. Asci emerging from the chlamydospore (asco-
genous cell) by rupture of the chlamydospore wall. (Fig. 20, D-G.)

Dimensiom: Of asci, 20-60[;, X 10-23ijl; of stalk cells, 7-20^^ X
10-30[x; of ascospores, 4-7[jl X 4-7[jl.

Distribution: Europe.

Material examined: Germany: Berlin, Aug., 1891, P. Sydow
(Myc. March. 3757, host given as A. hybrida but evidently A.
incana); Hamburg, Eppendorfer Moor, July, 1930, A. J. M. (two
collections) ; Rhine province, Westerwald, Stegskopf, July 30, 1922,
A. Ludwig (Mye. Ger. 1956). Norway: Orkedalen, July 16, 1887,
J. Brunchorst (Herb. Vaxtskyddsanstalt, Stockholm; host given
as A. glutinosa but evidently A. incana). Poland: Wilno, June
10, 1928, W.; Konopacka (Herb. Inst. Schol. Sup. Agric. Varsa-
viensis). Sweden: Jiimtland, Halland, July 23, 1915, 0. Juel;
Lappland, Abisko, Aug. 17, 1939, A. J. M.; ibid., Aug. 21, 1939, id.;
Stockholm, Bergianska Tradgarden, July 3, 1939, A. J. M. (col-
lections on A. incana, A. incana var. monstrosa Winkl., and var.
aurea Schelle.)

The following were collected as Taphrina klebahni: Germany:
Hamburg, Eppendorfer Moor, July, 1930, A. J. M. (two collections).
Norway: Ulvik, Aug. 18, 1935, A. J. M. Sweden: Stockholm, Ber-

Mix: Monograph of Genus Taphrina 85

gianska Tradgarden, Aug. 10-12, 1939, A. J. M. (collections on A.
incana, A. incana var. aurea and var. monstrosa) .

Wieben (1927) did not give dimensions of the stalk cells' of Taph-
rina klebahni, though from her description of the fungus it is clear
that she recognized their occurrence. The collections from Eppen-
dorfer Moor, Hamburg (the type-locality) were made with the
assistance of Miss Wieben.

Taphrina klebahni is evidently a late summer form of T. epi-
phylla. The first spots appear after the leaves of the witches' brooms
have borne asci and ascospores have been discharged. These spots
apparently result from infection by ascospores (or blastospores),
and presumably the resulting mycelium may overwinter and cause
a new witches' broom.

There is no morphological distinction between Taphrina epiphylla
and T. klebahni. Both have the same curious "tack-headed" asci
and the same pale-yellowish epiplasm. In size of asci, stalk cells,
and spores they are alike.

Both fungi seem to have the same biologic specialization. In the
Hortus Bergianus at Stockholm a plantation of Alnus was under
observation during the summer of 1939. Specimens of Alnus incana,
of A. incana var. aurea, and var. monstrosa, and of a variety with
pendulous branches, all showed witches' brooms of T. epiphylla
when observed early in July. In August when most of the leaves
had fallen from these witches' brooms, a heavy infestation of T.
klebahni appeared on the same trees. A specimen of A. incana var.
oxycanthoides Schotte, whose branches intermingled with a tree of
A. incana var. aurea remained free from both fungi, as did a nearby
tree of A. incana var. curvatipinnatifida Wittr. Specimens of Alnu^
rugosa (from North America) were heavily infested by Taphrina
sadebeckii and specimens of Alnv^ cordata Desf., and A. viridis DC
showed no signs of fungus attack.

The spots caused by Taphrina klebahni are usually small, one-
half to 1 cm. in diameter, but in the specimens collected at Ulvik,
Norway, all or nearly all of the leaf was involved, and only the ab-
sence of witches' brooms differentiated the fungus from typical
Taphrina epiphylla. Apparently the shoot-system was being at-
tacked by way of the leaves, and witches' brooms might be expected
to follow.

Wieben (1927) reported copulation of ascospores (or blasto-
spores) for both Taphrina epiphylla and T. klebahni. This phe-
nomenon has not been observed to occur regularly in any other
species.

86 The University Science Bulletin

49. Taphrina japonica Kusano

Taphrina japonica Kusano, Bot. Mag. Tokyo, 19:1-5. 1905, also Ann.

Mycol.' 3:30-31. 1905.

T. alni-japonicae Nishida, Miyabe Fertschrift. Tokyo, 1911.

T. macrophylla Ray, Mycologia 32:155-158. 1940.

Causing leaf curl with thickening, affecting part or all of the
blade, on leaves of Alnus japonica Sieb. and Zucc, and A. rubra
Bong.

Mycelium subcuticular.

Asci amphigenous, broad-cylindric, rounded at the apex, fre-
quently truncate at the base and broadened to a one-sided foot,
lacking a stalk cell. Ascospores eight, globose, frequently budding
in the ascus, filling it with ovate to elliptic blastospores (Fig. 21,
A-C.)

Dimensions: Of asci, 33-92[jl X 13-33[x, widening at base to 40[ji.
(if considered separately, T. japonica 46-92[jl X 23-33[jl, T. alni-
japonicae 36-85[x X 17-28[ji., T. macrophylla 36-86ii, X 13-26[x) , of as-
cospores, 4-5.5[J(,X4-5-5.5[ji,, of blastospores, 2-6.5[;,X2-5[ji,.

Distribution: Japan, Pacific coast of North America.

Material examined: Alnus japonica. Japan: Fukui, Tsudogama,
May 6, 1922, T. Asano; Ichikawa, June 9, 1907, S. Kusano; Iwate,
Morioka, June 21, 1904, G. Yamada (as T. alni-japonicae) ; ibid.,
July 26, 1905, id.; Tokyo, Botanic Garden, May 6, 1904, S. Kusano
(duplicate of type of T. japonica).

A. rubra. Alaska: June 26, 1923, J. P. Anderson. California:
Humboldt Co., Trinidad, March 24, 1931, H. E. Parks (Herb. Univ.
California 3592). Oregon: Tillamook Co., Siuslaw National For-
est, Hebo, May 9, 1923, J. S. Boyce (Herb. J. S. B. 1182).

Apparently Taphrina japonica Kusano, T. alni-japonicae Nish-
ida, and T. macrophylla Ray are one and the same fungus, for
which the name T. japonica has priority. Taphrina japonica is
said by Kusano to form witches' brooms, but judging from the way
this term is employed by many writers his statement may merely
mean that whole shoots and shoot-systems are affected. At any
rate the curled leaves produced by T. alni-japonicae may well be
due to infection by a fungus which will later induce a witches'
broom or deform a whole shoot (similar situations exist in Taphrina
cerasi and other witches'-broom formers). The leaves on the Jap-
anese specimens are not so greatly enlarged as in the American
specimens of the fungus on Alnus rubra. This may either be a host-
effect, or a difference that would disappear if more abundant Jap-
anese material were at hand.

Mix: Monograph of Genus Taphrina

fit

Fig. 21. A-C, asci (X 900) of Taphrina japonica, (B, as T. alni-japoni-
cae, C, as T. macrophylla) . D-H, T. occidentalis ; D, E, on Alnus rubra;
F, A. tenuifolia; G, A. sinuata; H, A. rhomhijolia.

88 The University Science Bulletin

The asci are alike in form and size. Measurements of the asci of
the three forms are given above. Asci of each fungus may widen at
the base to a broad diameter of 40[ji,.

Ray (1. c.) states that in Taphrina japonica "the basal portion
of the ascus is rounded and not widened as is often the case of the
asci on the leaves of A. rubra." Actually the asci of Taphrina jap-
onica, of T. alni-japonicae and of the fungus on Alnus rubra are
alike. They may be rounded at the base, or truncate and widened,
the widened portion often taking the form of a bilateral or unilat-
eral foot. Since there are no morphological differences between
these three fungi they are hereby declared synonymous as Taphrina
japonica Kusano.

The fungus deforming the leaves of Alnus rubra had been de-
scribed and discussed (Mix, 1939) before Ray's description ap-
peared, the question having been left open as to its identity with
the two Japanese fungi. The collection designated as the type of
T. macrophylla was not the earliest collection of the fungus, that be-
ing the one made by Boyce at Hebo, Oregon, May 9, 1923.

50. Taphrina occidentalis Ray
Taphrina occidentalis Ray, Mycologia 31 : 56-75. 1939.

Causing tongue-like enlargements of bracts of female catkins of
Alnus rhombifolia Nutt., A. rubra Bong, A. sinuata Ryd., and A.
tenuifolia Nutt. Also causing small puffed or curled areas on the
leaves of A. rhombifolia.

Mycelium subcuticular.

Asci covering surface of outgrowth, cylindric or cylindric-clavate,
rounded or truncate at the apex, with a stalk cell that is commonly
broader than long, truncate, rounded, pointed, or irregular below,
seated or somewhat inserted between epidermal cells. Ascospores
eight, round, ovate, or elliptic, frequently budding in the ascus, filling
it with smaller ovate or elliptic blastospores. Asci emerging from
the chlamydospore (ascogenous cell) by rupture of the chlamy-
dospore wall. (Fig. 21, D-H, Fig. 22, A.)

Dimensions: Of asci, 20-60ijl X 8-20iji,; of stalk cells, 5-23[jl X
l-23[).; of ascospores, 4-6[jl X 3.5-6iJL; of blastospores, S.5-5\i. X 3-4;;,.
Distribution: Western North America.

Material examined: Alnus rhombifolia. California: Lake Co.,
Putah Creek, Sept., 1931, H. E. Parks (Herb. Univ. Calif. 3883) ;
Monterey Co., Sur River, Aug., 1903, W. R. Dudley (Phanerogam.
Herb. Stanford Univ.) ; Santa Clara Co., Los Gatos Creek, Oct. 11,

Mix: Monograph of Genus Taphrina

89

Fig. 22. Asci (X900) of A, T. occidentalis on Alnus rhombifolia; B,
T. tosquinetii on A. glutinosa; C, A. crkpa var. mollis; D, the same
showing emergence of ascus from chlamydospore ; E, Taphrina viridis.

90 The University Science Bulletin

1902, W. R. Dudley (Phanerogam. Herb. Stanford Univ.). Ore-
gon: Prospect, Rogue River, Aug. 7, 1929, G. D. Darker (Herb.
Arnold Arboretum, 456) ; Sucker Creek below junction with Gray-
back Creek, Aug. 14, 1929, G. D. Darker (Herb. Arnold Arboretum
575).

A. rubra. British Columbia: D'Arcy, June 28, 1931, J. R.
Hansbrough (Herb. J. R. H. 605) ; Owl Creek, June 18, 1930, id.
(Herb. J. R. H. 606) ; Vancouver Island, Sooke, July, 1938, Malcolm
Wilson. California: Mill Valley, H. E. Parks (Fungi of Cali-
fornia 2596) ; Humboldt Co., Sur P. 0., Aug. 3, 1903, W. R. Dudley
(Phanerogam. Herb. Stanford Univ.) ; Siskiyou Co., Klamath Nat.
For., Deer Camp Ranger Station, Aug. 26, 1913, E. P. Meinecke
and J. S. Boyce (Herb. J. S. B. 278). Oregon: Corvallis, Jan. 30,
1938, Wm. Bridge Cooke; Grant's Pass, Sept. 5, 1917, J. R. Weir
(Herb. J. R. W. 5214. Type.)

A. sinuata. Oregon: Government Camp, July 23, 1929, G. D.
Darker (Herb. Arnold Arboretum 357) ; Oregon Caves, Lake Moun-
tain Trail, Aug. 12, 1929, G. D. Darker (Herb. Arnold Arboretum
532).

A. tenuifolia. Idaho: Clearwater Co., Elk River, Aug. 1929,
E. E. Hubert (Herb. J. S. B. 2073) ; Clearwater Nat. For., Bunga-
low Ranger Station, July 31, 1931, J. R. Hansbrough (Herb. J. R.
H. 607) ; Crystal Creek, Aug. 2, 1934, H. G. Lachmund and J. S.
Boyce (Herb. J. S. B. 2325). Montana: Glacier Nat. Park, Aug.,
1933, H. E. Bailey (Herb. Univ. California). Oregon: Oregon
Caves, Head of Limestone Creek, Aug. 15, 1929, G. D. Darker
(Herb. Arnold Arboretum 585). Washington: Walla Walla Co.,
Blue Mountain, Aug. 2, 1896, C. V. Piper (Washington Flora 401).

On leaves of Alnus rhombifolia. California: Mendocino Co.,
May, 1939, H. N. Hansen.

At the time of earlier publication (1939) this fungus was con-
sidered to be a variant of Taphrina robinsoniana. Ray (1939) has
since described it as a separate species. The asci of Taphrina occi-
dentalis are somewhat longer and wider, and the stalk cell is wider
than in T. robinsoniana. Taphrina occidentalis shows the further
peculiarity (not reported for T. robinsoniana) that the chlamy-
dospore wall breaks to allow emergence of the ascus.

The form on leaves of Alnus rhombifolia has stalk cells that are
slightly wider and more rounded than in any of the forms on cat-
kins. The ascus emerges from the chlamydospore in the manner
typical for this species. It is not impossible that this form on

Mix: Monograph of Genus Taphrina

91

leaves is a different species but for the time at least it seems best
to -include it in Taphrina occidentalis.

51. Taphrina viridis (Sadeb.) Maire

Taphrina viridis (Sadeb.) Maire, Bull. Soc. Bot. France 4e ser. 10:166-176.

1910.

Exoascus viridis Sadebeck (Jaap, Deutsch Bot. Monatschr. 19:75-76. 1901).

Taphrina alnastri Lagerheim, Vestergren Micr. Sel. Exs. 720. 1903.

Causing small, round, pale green or yellow spots (like those pro-
duced by T. sadebeckil) on leaves of Alnus viridis DC.

Mycelium intercellular.

Asci amphigenous, scattered, short-cylindric, or ellipsoidal-ob-
, long, rounded at the apex, stalk cells broad, rounded or truncate
below, inserted. Ascospores eight, budding in the ascus, ovate to
elliptic. Fig. 22, E.)

Dimensions: Of asci, 20-30[j. X T-lSpi.; of stalk cells, 7-13[jl X
8-18[x; of ascospores, 4.5-5[jl X 4-4. 5pi..

Distribution: Gennany, Austria (Brenner Post, Bad Ratzes),
Italian Alps, Scandinavia (according to Saccardo).

Material examined. Germany: Baden, Hollenthal, June 18, 1903,
G. Lagerheim (Micr. Sel. Exs. 720. Type of T. alnastri) ; ibid.,
July, 1903, id. (Bot. Mus. Stockholm).

This fungus is very similar to Taphrina sadebeckil but, as men-
tioned above, Alnus viridis is clearly not susceptible to attack by
the latter fungus, and it seems best to retain the two species as
distinct.

HOST INDEX TO SPECIES ON BETULACEAE

Alnus crispa (Ait.) Pursh. var. ynoUis Fern.

Alnus glutinosa Gaertn.

Alnus hirsuta Turcz.

Alnus hirsuta Turcz. var. sibirica Schneid.

Alnus hybrida A. Br.

Alnus incana Moench.

Alnus incana Moench. var. aurea Schelle

Alnus incana Moench. var. monstrosa Winkl.

Alnus iaponica 8ieb. and Zucc.

.'ilnus rhombifolia Nutt.

Alnus rubra Bong.

Alnus rugosa Spreng.

Alnus sinuata Rydb.
Alnus tenuifolia Nutt.

Taphrina tosquinetii (Westend
Taphrina amentorum (Sadeb.)
Taphrina sadebeckii Johans.
Taphrina tosquinetii (Westend
Taphrina amentorum (Sadeb.)
Taphrina sadebeckii Johans.
Taphrina sadebeckii Johans.
Taphrina amentorum (Sadeb.)
Taphrina tosquinetii (Westend.
Taphrina amentorum. (Sadeb.)
Taphrina epiphylla Sadeb.
Taphrina robinsoniana Gies.
Taphrina epiphylla Sadeb.
Taphrina epiphylla Sadeb.
Taphrina japonica Kus.
Taphrina occidentalis Ray
Taphrina amentoruTn (Sadeb.)
Taphrina japonica Kus.
Taphrina occidentalis Ray
Taphrina robinsoniana Gies.
Taphrina sadebeckii Johans.
Taphrina occidentalis Ray
Taphrina occidentalis Ray

) Tul.
Rostr.

) Tul.

Rostr.

Rostr.
) Tul.
Rostr.

Rostr.

92

The University Science Bulletin

Alnus viridis DC.
Betula aurata Bechst.
Betula carpatica Waldst.

Betula ermani Cham.

Betula fontinalis Sarg.
Betula fruticosa Pall.
Betula glandulosa Michx.
Betula huviilis Schrank.

Betula intermedia Thom.

Betula japonica Sieb.
Betula lutea Michx.
Betula medwediewi Reg.

Betula nana L.

Betula oceidentalis Hook.
Betula papyrifera Marsh.

Betula pendula Roth.
Betula populifolia Ait.

Betula pubescens Ehrh.

Carpinus betulus L.
Carpinus caroliniana Walt.
Carpinus orientalis Mill.
Corylus americana Marsh.
Corylus heterophylla Fisch.
Corylus rostrata Ait.
Corylus sieboldiana Blume
Ostrya carpinifolia Scop.
Ostrya virginica Willd.

TaplLrina

Taphrina

Taphrina

I Taphrina

1 Taphrina

I Taphrina

"1 Taphrina

Taphrina

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Taphrina

viridis Maire
betulina Rostr.
betulina Rostr.
betuUcola Nish
nana Johans.
americana Mix
boycei Mix
carnea Johans.
bacteriosperma
carnea Johans.
carnea Johans.
bacteriosperma
betulae (Fkl.)
betulina Rostr.
carnea Johans.
nana Johans.
nana Johans.
americana Mix
carnea Johans.
betulae (Fkl.)
bacteriosperma
betulina Rristr.
carnea Johans.
nana Johans.
boycei Mix
americana Mix
carnea Johans.
flava Farl.
betulae (Fkl.)
betulina Rostr.
carnea Johans.
nana Johans.
flava Farl.
bacteriosperma
betulae (Fkl.)
betulina Rostr.
carnea Johans.
nana Johans.
carpini (Rostr.
australis (Atk
carpini (Rostr.

Johans.

Johans.
Johans.

Johans.
Johans.

Johans.

Johans.
Johans.

) Johans.
) Gies.
) Johans.

Taphrina coryli Nish.

Taphrina ostryae Massal.
Taphrina virginica Sadeb.

Mix: Monograph of Genus Taphrina 93

IV. SPECIES ON FAGACEAE

Fagus, Nothofagus, Castanopsis, Quercus

Laubert (1928) lists Taphrina fagi Lamb., as occurring on Fagus
sp. in North America. No specimen of such a fungus could be found
in the Berlin Museum, whose collection Laubert used in writing his
account of known species. When appealed to by letter in 1938 he
was unable to recall the source of his information. Thorough search
of the literature has yielded no other reference to such a fungus.

52. Taphrina entomospora Thaxter

Taphrina entomospora Thaxter, Bot. Gaz. 50:435. 1910.
Exoascus entomosporus (Thaxt.) Saccardo and Trotter, Sylloge Fungorum
22:765.

Entomospora antarctica (Sacc.) Jaczewski, Pocket key for the determina-
tion of fungi. Part I. Exoascales. Leningrad. 1926.

Causing yellowing (without enlargement or thickening) of leaves
of Nothofagus antarctica Oerst. All the leaves of a shoot may be
attacked.

Mycelium subcuticular.

Asci hypophyllous, subcylindric, rounded or subtruncate at the
apex, provided with a stalk cell which is usually broader than the
ascus. Ascospores eight, described by Thaxter as bearing appen-
dages of two orders.

Dimensions: Of asci, 36-53[x X 10-17[jl; of stalk cells, 10-13[x X
17-23[x (Thaxter reports asci 55-60[ji X 13-15[jl, perhaps including
the stalk cell in his measurements) ; of ascospores, 3.5-6[ji, X 2-4. 5^/,;
of blastospores, 10-23ijl X 0.5-1. Opi. (Thaxter reports ascospores as
9-10[x X 3-4[jL, with terminal appendages 8-12[jl X S-5\k, subterminal
appendages 15-25[ji, X 0.8[;..) (Fig. 23, A.)

Distribution: Patagonia.

Material examined: Chile: Puntas Arenas, Feb., 1906, R. Thax-
ter (Type).

Perhaps the portion of material examined was not so favorable as
that studied by Thaxter. Many asci were old and empty. Only a
few appendaged spores were seen, and these were not so beautifully
regular as those seen by Thaxter. The appendages looked like nar-
row elongate buds, and many asci contained numerous filiform blas-
tospores. Most ascospores observed were elliptic, and either lacked
appendages or bore one or more short, narrow buds.

94

The University Science Bulletin

Fig. 23. Asci of A, Taphrina entomospora ; B, T. kusanoi; C, T.
castanopsidis. D, outlines of asci and ascus-bases of T. caerulescens on
Qucrnis gamhclii; E, Q. mongolica; F, Q. prinoides; G, Q. utahensis.
All X 900.

Mix: Monograph of Genus Taphrina 95

53. Taphrina castanopsidis Jenkins
Taphrina castanopsidis Jenkins, Mycologia 28:31-34. 1936.

Causing convex-concave spots on leaves of Castanopsis chryso-
phylla DC.

Mycelium intercellular.

Asci very long, cylindric, narrow, rounded at the apex, narrowed
at the base into one or more short rhizoidal extensions, inserted be-
tween epidermal cells, no stalk cell present. Ascospores round,
eight, budding in the ascus and filling it with smaller ovate to ellip-
tic blastospores.

Dimensions: Of asci, 80-165[jl X 10-17[x; of ascospores, diameter
up to 10[ji (according to Jenkins) ; of blastospores, 3-5[l X 1.5-2.5pL.
(Fig. 23, C.)

Distribution: California.

Material examined: California: Calaveras Co., Dorrington,
Aug. 16, 1934, J. S. Boyce (Herb. J. S. B. 2287) ; region of the upper
Sacramento, July 24-Aug. 10, 1894, Sisson (Plants Univ. Calif., 121
Fungi of Calif, as T. castanicola E. and E. n. sp.) ; Siskiyou Co., Mt.
Shasta, Aug. 30, 1937, Wm. Bridge Cooke.

54. Taphrina kusanoi Ikeno
Taphrina kusanoi Ikeno, Flora 92:1-31. 1903.

Causing small (up to 1 cm. diam.) convex-concave spots on leaves
of Castanopsis cuspidata Schottky {Pasania cuspidata Oerst.).

Mycelium intercellular.

Asci hypophyllous, cylindric, rounded at the apex, often widened
at the base, or provided with a distinct foot, lacking a stalk cell.
Ascospores not observed, asci filled with round, ovate, or elliptic
blastospores, some of them very small. (Fig. 23, B.)

Dimensions: Of asci, 36-80[x X 12-20[a (102-117iji. X 13-19[ji, ac-
cording to Ikeno) ; of blastospores, 2-5[k X 2-3ijl or very minute.

Distribution: Japan.

Material examined: Japan: Idzen, Atanei, April 13, 1895; Mt.
Tsukuba, May, 1900, Kusano; ibid., June 19, 1929, id.; Tokyo,
Saginomiya, June 19, 1903, Y. Shibasaksi (Herb. Morioka Imp.
Coll. Agric. and For.).

If this species actually possesses asci of the size given by Ikeno,
the possibility of its synonymy with Taphrina castanopsidis may
arise. However, in specimens examined the asci were definitely
shorter than those of T. castanopsidis and the expanded foot of the
ascus is a distinguishing feature.

96

The University Science Bulletin

Qimbricdrid O.Pdlusiri5 0 lobatd Omaximd OMirbecKii

Fig. 24. Asci and ascus-bases (X 900) of Taphrina caerulescens, on

species of Quercus.

Mix: Monograph of Genus Taphrina 97

55. Taphrina caerulescens (Desm.) Tulasne

Taphrina caerulescens (Desm.) Tulasne, Ann. Sci. Nat. 5e Ser. Bot. 5:122-

136. 1866.

Ascomyces caerulescens Desmazieres, Ann. Sci. Nat. 3e Scr. Bot. 10:342-361.

1848.

A. alutaceus Thiimen, Verhandl. K. K. Zool. Bot. Ges. Wien. 29:523-524.

1880.

A. extensus Peck, New York State Mus. Nat. Hist. Ann. Rept. 39:30-70.

1886.

A. ruhrohrunneus Peck, New York State Mus. Nat. Hist. Ann. Rept. 40:39-

77. 1887.

Causing small to large, convex-concave, slightly swollen spots or
blisters on leaves (occasionally deforming the whole leaf, but not
forming true witches' brooms) on many species of Quercus. (A
complete list, as far as known, is given below.)

Mycelium intercellular.

Asci epiphyllous or hypophyllous, very variable in size and shape,
cylindric or clavate, usually rounded at the apex, at the base blunt,
rounded, or truncate, and seated or slightly inserted; or else wedge-
shaped, pointed, or with rhizoidal appendages, and more or less
deeply inserted between epidermal cells. Stalk cells lacking. Asco-
spores rarely seen, asci filled with round, ovate, or elliptic blasto-
spores. (Fig. 23, D-G, Fig. 24, 25.)

Dimensions of asci in microns

Quercus acutissima Carruth 50- 92 X 15-27

Quercus agrijolia Nee 46- 68 X 15-19

Quercus alba L. 49-103 X 15-34

Quercus borealis Michx 46- 76 X 15-30

Quercus cerris L 30- 72 X 11-23

Quercus cinerea Michx 42- 68 X 15-23

Quercus coccijera L 46- 84 X 11-27

Quercus coccinea Muench 68-118 X 13-30

Quercus crispula Blume 46- 92 X 14-23

Quercus dentata Thunb 57- 84 X 13-22

Quercus douglasii Hook, and Am 65-118 X 19-28

Quercus ellipsoidalis Hill 57-104 X 15-30

Quercus jruticosa Brot 38- 65 X 11-19

Quercus gambelii Nutt 53- 83 X 20-27

Quercus geminala Small 75-120 X 15-27

Quercus georgiana Curtis 49- 84 X 19-30

Quercus ilicifolia Wangh 46- 84 X 15-27

Quercus imbricaria Michx 49- 91 X 11-30

Quercus kelloggii Newb 65-106 X 19-27

Quercus laurifolia Michx 57- 95 X 15-27

Quercus lobata Nee 50- 87 X 17-23

Quercus macrocarpa Michx 38- 76 X 15-27

Quercus marilandica Muench 53-103 X 11-26

Quercus maxima Ashe 46- 95 X 13-30

Quercus mirbeckii Dur 46- 84 X 19-27

Quercus mongolica Turcz 76-109 X 20-26

Ouercus nigra L 60-102 X 19-34

Quercus palustris Moench 58- 84 X 19-27

7—3659

98 The University Science Bulletin

Quercus phellos L 46- 76 X 11-19

Quercus prinoides L 32- 58 X 18-22

Quercus pubescens Willd. (Q. lanuginose) 43- 84 X 15-27

Quercus rubra L 57- 99 X 17-25

Quercus robur L 50- 84 X 19-25

Quercus serrata Thunb 64- 92 X 18-30

Quercus sessiliflora Salisb 46- 72 X 15-27

Quercus stellata Wangh 57- 80 X 15-27

Quercus undulata Torr 38- 84 X 15-27

Quercus utahensis Rydb 40- 73 X 20-27

Quercus velutina Lam 49- 95 X 15-27

Quercus virginiana Mill 68- 95 X 15-23

Ascus-size limits for all known host-species: 30-120[x X ll-34[ji,.

Material examined: Quercus acutissima. Japan: Iwate, Mizu-
yama, July 25, 1927, K. Togashi (Herb. K. T. 2899) ; ibid., June 16,
1932, id. (Herb. Morioka Imp. Coll. Agric. and For.).

Q. agrijolia. California: Monterey Co., April, 1938, H. Earl
Thomas; ibid., June 16, 1932, H. N. Hansen.

Q. alba. Massachusetts: Waltham, June 20, 1935, D. H. Lin-
der. New York: Ithaca, 1938, W. W. Ray. Virginia: Victoria,
June 9, 1936, S. A. Wingard.

Q. borealis. Connecticut: East Granby, July 4, 1934, H. G.
Eno (U. S. D.A. For. Path. 81724). Massachusetts: Petersham,
July 26, 1928, J. S. Boyce (Herb. J. S. B. 2028). Michigan: Huron
Co., Sand Point, near Bayport, July 4, 1938, E. A. Bessey. New
Hampshire: Conway, June 8, 1921, P. Spaulding (For. Path.
45371). Tennessee: Great Smoky Mts., Nat. Park, Rich Mt.,
June 24, 1934, L. R. Hesler.

Q. cerris. Algier: Guide, Kofouri, Sept. 16, R. Maire and M.
Pettitmengin (Miss. Bot. Or. 1906). Italy: Mt. Penice, Vocarezza
(Cav. F. Longob. Exs. 73a) ; Ponte Organasco (Cav. F. Longob.
Exs. 73b).

Q. cinerea. North Carolina: Aiken, 1878, H. W. Ravenel
(Thum, Myc. Univ. 2065).

Q. coccifera. Algier: Marabout de Sidi-Youssef, Bouzaria, May
8, 1913, R. Maire (Myc. Bor. Afr.). France: Hyeres, June 16
(Rab.-Wint. F. Eur. 3537). Greece: Mt Ypsili-Keryphi, near
monasteiy Renive, July 12, 1906, R. Maire (Miss. Bot. Or. 240).
Morocco: Cap Spartel, April 26, 1924, R. Maire (Champ. Afr.
Nord. 8403).

Q. coccinea. Georgia: Athens, May 14, 1936, J. H. Miller.
Michigan: Huron Co., Sand Point, near Bayport, July 4, 1938, E.
A. Bessey. New York: Ithaca, South Hill Marsh, June 26, 1940,
A.J.M.

Mix: Monograph of Genus Taphrina

99

Q.$emincil(i Q. Kfll068ii O-vir^miand Oldurifolid

Q.d?ntflld O-slelldld Q.robur Ophellos 0.5e55iliflord

0-undulaia Oasnfolia Qc^lba 0. ciner^d 0-trUlicosd

Fid 25. Asci and ascus bases (X900) of Taphrina caerulescens, on

species of Quercus.

100 The University Science Bulletin

Q. dentata. Saskatchewan: Indian Head, July 25, 1935, B. J.
Sollano (Herb. Univ. Toronto).

Q. douglasii. California: Lake Co., May 3, 1936, H. N. Han-
sen.

Q. ellipsoidalis. Wisconsin: Lyndon Station, July 4, 1917, J. J.
Davis (Herb. Univ. Wisconsin) .

Q. fruticosa. Lusitania: Coimbra, summer 1879, A. F. Moller
(Myc. Univ. 1553).

Q. gamhelii. Wyoming: Medicine Bow Nat. For., Hayden Divn.,
Sandstone Ranger Sta., July 13, 1939, W. G. Solheim.

Q. geminata. North Carolina: May, 1936, R. F. Poole.

Q. georgiana. Georgia: Stone Mt., April 26, 1925, J. H. Miller
(Herb. Univ. Georgia).

Q. ilicifolia. New Hampshire: W. Ossipee, July 19, 1910, W. G.
Farlow (Farl. Herb.). New York: Sam's Point, Aug., J. Dearness
(Herb. J. D.). Virginia: Bald Knob, July, 1918, G. H. Chapman
(Herb. Mass. Agric. Coll. 2486).

Q. imbricaria. Missouri: Perry vale, C. H. Demetrio (Rab-
Pazsch. F. Eur.). North Carolina: Greensboro, June, 1934, A. F.
Thiel ; ibid., spring 1934, E. M. Martin.

Q. kelloggii. California: Lake Co., May 3, 1936, H. N. Hansen.

Q. laurifolia. Florida: Gainesville, May 20, 1935, G. F. Weber;
ibid., April 28, 1941, id.

Q. lobata. California: 1935, H. N. Hansen.

Q. macrocarpa. Alberta: J. Dearness (Herb. J. D.). Iowa:
W. Okeboji, June 19, 1932, G. W. Martin. Kansas: Hays, June
6, 1930, E. Bartholomew (N. A. F. 10957, as T. externa) ; Lawrence,
Haskell Meadow, May 14, 1946, A. J. M. Wisconsin: Granville,
July 17, 1867, I. A. Lapham.

Q. marilandica. Kansas: Baldwin, June, 1933, A. J. M.; ibid.,
June 2, 19S6, id.; ibid.. May 31, 1937, id. Missouri: Bagnell Dam,
Osage Beach, June 3, 1937, A. J. M.; Seligman, July 12, 1922, R. P.
White. Virginia: Princess Anne Co., May 18, 1926, H. T. Cook.

Q. maxima. Connecticut: Collinsville, Aug., 1937, A. J. M.;
New Haven, June 23, 1928, id.; ibid., June 30, 1928, id. Georgia:
Athens, May 14, 1936, J. H. Miller; Experiment, May 18, 1936, J. L.
Weimer. Maine: Kittery Point, June, 1899, R. Thaxter (Farl.
Plerb.). Massachusetts: Amherst, Aug. 19, 1937, A. J. M.; Man-
chester, Sept. 4, 1928, R. J. Eaton (For. Path. 45878) ; Sunderland,
Aug., 1934, R. H. Thompson. New Hampshire: Canaan, Sept. 23,
1931, E.D. Farnsworth (For. Path. 51682). New York: Columbia
Co., near Hudson, Aug., 1919, A. J. M.; Ithaca: Fall Creek Ravine,

Mix: Monograph of Genus Taphrina 101

near Beebe Lake, June 27, 1940, id.; Poultry Woodlot, Aug. 18,
1929, D. S. Welch (Herb. D. S. W. 819) ; Klinewoods Road, June 21,
1940, A. J. M.; Sandlake, Sept., 1886, C. H. Peck [Type of T. rubro-
brunnea). Ontario: Muskoka, Muldrew Lake, July 31, 1936, D. S.
Welch (Herb. D. S. W. 1286). North Carolina: 1937, J. N. Couch
(Herb. Univ. North Carolina, 10724). Tennessee, June 20, 1937,
C. D. Sherbakoff. Wisconsin: June 2, 1913, J. J. Davis (Herb.
Univ. Wisconsin).

Q. mirbeckii. Morocco: Janger a Agla, April 25, 1924, R. Maire
(Champ. Afr. Nord. 8404) .

Q. mongolica. Russia: Nyushno-Yssyriiskii Krai, June 13, 1912,
N. Naumov (Herb. Sydow).

Q. nigra. Alabama: Auburn, May 8, 1890, G. F. Atkinson
(Econ. F. 180). Florida: Gainesville, April 28, 1941, G. F. Weber.
Massachusetts: Melrose, June 16, 1936, R. H. Thompson. North
Carolina: Durham, F. A. Wolf (Herb. Univ. Tennessee 9978).
Q. palustris. Georgia: Athens, May 14, 1936, J. H. Miller.
Q. pedunculata. Germany: Schleswig-Holstein, Island Sylt,
Lornsenhain, near Westerland, May 8, 1911, H. and P. Sydow
(Myc. Germ.).

Q. phellos. Alabama: Auburn, May 8, 1890, G. F. Atkinson.
North Carolina: 1938, F. A. Wolf.

Q. prinoides. Kansas: Vinland, Violet Hill, May 14, 1946. A.
J. Mix.

Q. pubescens. Austria: Kaltenleutgeben, Aug., 1879, Thiimen
(Myc. Univ. 1554, as Ascomyces alutaceus). Italy: Verona, Tre-
gagno, autumn 1906, C. Massalongo (Myc. Ital. Exs. 1678).
Q. robur. Ireland: Glengariff, June 18, 1935, A. J. M.
Q. rubra. Alabama: Auburn, May 13, 1890, G. F. Atkinson
(Econ. F. 185). North Carolina: Durham, June, 1935, F. A.
Wolf.

Q. sessiliflora. Italy: Florence, Vallombrosa, spring 1898, F.
Cavara (Myc. Ital. 534). Montenegro: Savnitz, Sept. 30, 1911, E.
Vlack (BubakHerb.).

Q. stellata. Illinois: Mount Vernon, Aug. 6, 1937, G. H. Boewe.
South Carolina: Clemson College, June 15, 1935.

Q. undulata. Colorado: Mancos, June 22, 1898, Baker, Earle
and Tracy (Farl. Herb.) ; Ouray, July 4, 1907, Clements (Crypt.
Form Colo. 527).

Q. utahensis. Colorado: Glenwood Springs, Aug. 20, 1941, A.
J.M.

102 The University Science Bulletin

Q. velutina. Arkansas: Fayetteville, May 21, 1935, J. C. Dune-
gan. Connecticut: Collinsville, Aug. 17, 1937, A. J. M.; New
Haven, June 24, 1928, id.; New London, June, 1886, W. G. Farlow
(Farl. Herb.). Massachusetts: Hamilton, July, 1927, P. Spauld-^^
ing (For Path. 16130) ; Middlesex Fells, A. B. Seymour; Pigeon^
Cove, July 28, 1890, id. (Econ. F. 184b). Mississippi: Starkville,
May 10, 1890, S. M. Tracy (Econ. F. 184a). Missouri: Camden-
ton, June 3, 1937, A. J. M. West Virginia: Monongahela Co., near
Lake Lynn, July 12, 1935, C. R. Orton. Wisconsin: Avoca, July
18, 1923, J. J. Davis (Herb. Univ. Wisconsin).

Q. virginiana. Virginia: Princess Anne Co., May 21, 1935, H. T.
Cook; ibid., May 18, 1936, ^d.

Besides the hosts listed above T. caendescens is reported on:
Quercus armeniaca Kotsch in the Caucasus (Jacsewski) ; Q. aus-
trina Poplarville, Mississippi, Aug. 7, 1925, J. L. Weimer; Quercus
conferta Kit., Romania (letter from Savelescu, 1939) ; Q. gunnisonii,
North Cheyenne Canyon, Colorado, July, 1895, L. H. Pammel.

The above account of Taphrina caerulescens is taken from the
paper by Mrs. Thompson (1940) with a few emendations and
studies of additional species by the writer, and Figures 24 and 25 are
copied from her illustrations.

Since the chief point of distinction between Taphrina caerulescens
and T. kruchii (see below) lies in the fact that the latter forms
witches' brooms, a collection in the Farlow Herbarium becomes in-
teresting. It is labelled: "Taphrina on red oak, Kittery Point,
Maine, June, 1899, Thaxter." Enclosed is a slip of paper with the
notation, apparently by Thaxter: "T. kruchii."

The specimens show clusters of diseased leaves arising in some
cases from the ends of twigs, in some cases from axillary buds. All
leaves are reduced in size and deformed throughout the whole blade.
No adventitious buds are present and in spite of their suggestive
appearance the structures are not true witches' brooms. The asci
conform to those of Taphrina caerulescens on Quercus maxima.

56. Taphrina kruchii (Vuill.) Saccardo

Taphrina kruchii (Vuill.) Saccardo, Sylloge Fungorum 10:68.
Exoascus kruchii Vuillemin, Rev. Mycol. 13:141-142. 1892.

Causing pronounced witches' brooms with negatively geotropic
curvature of twigs, on Quercus ilex L.
Mycelium intercellular.
Asci hypophyllous, cylindric to clavate, rounded at the apex,

Mix: Monograph of Genus Taphrina

103

Fig. 26. Asci (X900) of A, Taphrina kruchii; B, T. ulmi ; C, T. celtis
on Celtis anstralis; D, C. toumejortii; E, Taphrina bullata.

104 The University Science Bulletin

without stalk cell. Ascospores eight, asci usually filled with elliptic
or rod-shaped blastospores. (Fig. 26, A.)

Dimensions: Of asci, lO-lOOpL X 15-25[jl; of ascospores, 4\i diam.
(Vuillemin) ; of blastospores, 2.5^ X Spi. (Vuillemin) ; of rod-shaped
blastospores, S-IO^jl X l-5-2\i.

Distribution: Mediterranean area.

Material examined: Algier: between Babor and Tababor, hill
"Tizi n' Souk," May 7, 1912, R. Maire (Myc. Bor. Afr. 98, asci not
found). Corsica: Zuza, Sept. 6, 1901, R. Maire (Micr. Rar. Sel.
662). Italy: Cagliari, Sette Fratella, July, 1900, Cavara (Myc.
It. 687). Sardinia: Mt. Gennargentii, Aug., 1902, A. Firoi (Myc.
It. 1517). Sicily: Albano, June 2, 1914, P. Baccoirini (Herb. R.
Mus. Florence).

This fungus, whose asci conform to those of Taphrina caeniles-
cens, is retained as a separate species because of its habit of forming
witches' brooms, and its probable (though unknown) biological
specialization.

The specimen (mentioned above) from Albano, Sicily, obtained
on loan from the herbarium of the Royal Museum, Florence, is of a
large witches' broom with many crowded adventitious shoots. The
twigs show negative geotropic curvature, and the leaves are dwarfed
and completely deformed by the fungus.

HOST INDEX TO SPECIES OF TAPHRINA ON FAGACEAE

(Exclusive of Taphrina caerulescens)

Castanopsis chrysophylla DC

Taphrina castanopsidis Jenkins
Castanopsis ciispidata Schottky

Taphrina kusanoi Ikeno
Nothojagus antarctica Oerst.

Taphrina entomospora Thaxt.
Quercus ilex L.

Taphrina kruchii Vuill.

Mix: Monograph of Genus Taphrina 105

IV. SPECIES ON URTICACEAE

Ulmus, Celtis
57. Taphrina ulmi (Fkl.) Johanson

Taphrina ulmi (Fkl.) Johanson, Ofvers K. Svensk. Vet. Akad. Forhandl.

1885:20-47. 1886.

Exoascus ulmi Fuckel, Jahrb. Nassau. Verein. Naturk. 27 and 28:1-99.

1873 and 1874.

Exoascus campestris Saccardo, Michelia 2:30-135. 1892.

Causing small, yellowish to brown, unthickened or slightly puffed
spots on leaves of Ulmus alata Michx., U. americana L., U. fulva
Miehx., U. glabra Huds., U. laevis Pall., U. montana With., U.
carpinifolia Gleditsch var. suberosa (Moench.) Rehd.

Mycelium subcuticular.

Asci hypophyllous or sometimes epiphyllous, cylindric or ellipsoid,
rounded at apex, with a broad stalk cell. Ascospores eight, round,
ovate, or elliptic, often budding in the ascus. (Fig. 26, B.)

Dimensions: Of asci, 10-29[jl X 7-15[x; of stalk cells, G-lOpi. X
8-22[j,; of ascospores, S-6\>. X 3-6[x.

Distribution: Europe, North America.

Material examined: Ulmus alata. Alabama: Tuskegee, March
29, 1897, G. W. Carver (Ellis Herb.). Arkansas: near Fayetteville,
May, 1941, received by J. C. Dunegan.

U. americana. Quebec: Duchesnay, 1935, R. Pomerleau; ibid.,
June 12, 1941, id.

U. julva. Kansas: Baldwin, June 15, 1947, A. J. M. Virginia:
Blacksburg, June, 1932, S. A. Wingard (U. S. D. A. Path, and Myc.
Coll. 66890).

U. glabra. England: Kent, Ivy Hatch, July 10, 1935, A. J. M.
Poland: Pulawy, May 20, 1927, W. Siemaszko (Herb. Inst. Phyto-
path. Schol. Sup. Varsaviensis) . Sweden: Uppsala, June, 1885, C. J.
Johanson; Skane, Pelshult, Aug., 1896, G. Lagerheim.

U. laevis. Germany: Saxony, between Schmilka and Herrnskret-
chen. May 26, 1899, W. Krieger (F. Sax. 622). Holland: Peters-
berg, near Maastrecht, Aug., 1885, P. Magnus (Rehm. Asco. 869) .

106 The University Science Bulletin

58. Taphrina celtis Sadebeck

Taphrina celtis Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 8:61-95. 1890.

Exoascus celtidis (Sadeb.) Saccardo, Sylloge Fungorum 10:69.

E. aemiliae Passer., Atti. R. Accad. Lincei, Ser. 4, 6:457-470. "1889" (1890).

Causing small roundish, gray to brown, very slightly thickened
spots, apparent on upper surface only, on leaves of Celtis australis
L., and C. tournefortii Lam.

Mycelium subcuticular.

Asci hypophyllous, cylindric to ellipsoid, rounded at the apex;
stalk cell flattened, sometimes broader than the ascus. Ascospores
eight, round, ovate, or elliptic, often budding in the ascus. (Fig.
26, C, D.)

Dimensions: Of asci, 13-28[jl X 8-13[;,; of stalk cells, 7-10[;. X
8-13[i, (8-10[jL X 25-30[jL according to Sadebeck) ; of ascospores,
3.5-5[x X 3.5-4.51;,.

Distribution: southern Europe, north Africa,

Material examined. Celtis australis. Algier: Sidi-Madani,
Gorges de la Cluffa, May 4, 1913, R. Maire (Myc. Bor. Afr. 170).
Caucasus: Tiflis, Hortus Botanicus, 1911, G. Nevodowski.

C. tournefortii. Herzegovina: Trebinje, May 9, 1897, C. Barinitz
(Herb. Sydow),

HOST LIST FOR TAPHRINA ULMI (FKL.) JOHANS

Ulmus alata Michx.

Ulmus americana L.

Ulmus carpinifolia Gleditsch var. subernsa (Mocncli.) Rohrd

Ulmus fulva Michx.

Ulmus glabra Huds.

Ulm,us laevis Pall.

Ulmus montana With.

Mix: Monograph of Genus Taphrina 107

V. SPECIES ON ROSACEAE

Pyrus, Sorbus, A^nelanchier , Crataegus, Potentilla, Geum

59. Taphrina bullata (Berk.) Tulasne

Taphrina bullata (Berk.) Tulasne, Ann. Sci. Nat., 5 Ser., Bot. 5:122-136.

1866.

Ascomyces bullatus Berkeley, Jour. Roy. Hort. Soc. London. 9:48. 1854.

Exoascus bullatus (Berk.) Sadebeck, Jahrb. Hamburg. Wissensch. Anst.

10:5-110. 1893.

Causing small, slightly thickened, bullate, irregular brown spots on
leaves of Pyrus communis L.

Mycelium subcuticular.

Asci hypophyllous, cylindric, rounded or truncate at the apex,
stalk cell as broad as, or narrower than the ascus. Ascospores eight,
round, ovate, or elliptic, often budding in the ascus (Fig. 26, E.)

Dimensions: Of asci, 23-36tJi, X 8-15[jl (30-40[i, X S-Qpt. according
to Sadebeck) ; of stalk cells, 8-17[jl X 5-13[x; of ascospores, 4.5-5.5[x
X 3.5-4.5[x.

Distribution: Europe.

Material examined: Germany: Berlin, Steglitz, Aug. 18, 1887, P.
Magnus; Brandenburg, Tamsel Baumschulen, Aug. 26, 1904, P.
Vogel (Myc. Germ. 391). Poland: Zalesczyki, May 25, 1910, A.
Wroblewski (Myc. Polon. II 54). Sweden: Uppsala: Bot. Gard.,
June 8, 1912, 0. Juel; Stockholm, Experimentalfaltet, June 28, 1939,
A. J. M.; ibid., July 25, 1939, id.; Vaxtskyddsanstalt, July 1, 1939,
id.; Tungelsta, July 13, 1939, id.

Buhr (1935) reports having seen in the Bremen Botanic Garden
Taphrina bullata on Pyrus betulijolia Bunge, P. sinensis Lindl., and
"P. sinensis X salici folia X communis."

Pyrus sinensis Lindl. is P. lindleyi Rehd. and on this host the writer
(1947) has described a new species Taphrina orientalis (see below).
Buhr (1. c.) also states that T. bullata occurs at Corte' in Corsica on
P. amygdaliformis Vill. The fungus is also reported by Tai (1937)
as occurring on P. montana Nakai, at Port Arthur, Dairen. The re-
port by Rostrup (1890) of Taphrina bullata as occurring at Viborg,
Denmark, on Chaenomeles lagenaria Koidz. {Cydonia japonica) is
in error. This specimen (obtained on loan from the Copenhagen
Museum) is plainly Pyrus communis, though perhaps a seedling
rather than a cultivated variety.

108

The University Science Bulletin

Fig. 27. Asci (X 900) of A, Taphrina orientalis; B, T. piri; C, T. sorbi;
D, T. amelanchieri; E, T. cralaegi; F, T. potentillae.

Mix: Monograph of Genus Taphrina 109

60. Taphrina orientalis Mix
Taphrina orientalis Mix, Trans. Kansas Acad. Sci. 50:77-83. 1947.

Causing small (up to 8 mm. diam.) deformed spots on leaves of
Pyrus lindleyi Rehd.

Mycelium subcuticular.

Asci hypophyllous, cylindric, lacking a stalk cell, seated on the
epidermis. Ascospores eight, frequently budding and filling the
ascus with blastospores. (Fig. 27, A.)

Dimensions: Of asci, 22-46[x X 8-17tx; of ascospores, 4-6[jl X 4-5[i..

Distribution: Japan.

Material examined: Japan: Pref. Iwate, Morioka, May 30, 1908,
G. Yamada (Received from K. Togashi.) .

61. Taphrina piri Kusano

Taphrina piri Kusano, Bot. Mag. Tokyo 19:1-3. 1905, and Ann. Mycol.

3:30-31. 1905.

Exoascus piri (Kus.) Saccardo, Sylloge Fungorum 18:197.

Causing small, pale, bullate spots or larger curled areas on leaves
of Sorbus alnifolia K. Koch. {Pyrus miyabei Sarg.).

Mycelium subcuticular.

Asci hypophyllous, cylindric, rounded or truncate at the apex,
occasionally broadened to a foot below; lacking stalk cells, as-
cospores eight, round, ovate, or elliptic, frequently budding in the
ascus. (Fig. 27, B..)

Dimensions: Of asci, 23-42[jl X 8-13[jl; of ascospores, 4:-6[i X
3.5-51JL.

Distribution: Japan.

Material examined: Japan: Nikko, June 10, 1904, S. Kusano

(duphcate of type, received from Kusano) ; ibid., id. (in Bot. Mus.

Stockholm) .

62. Taphrina sorbi (Jacz.) Mix

Taphrina sorbi (Jacz.) Mix, Univ. Kansas Sci. Bull. 24:10:151-176. 1936.
Exoascus sorbi Jaczewski, Pocket Key for Determination of Fungi. Part
I Exoascales. Leningrad. 1926.

Causing small deformed spots, or leaf-curl in leaves of Sorbus
torminalis' (L.) Crantz.

Mycelium subcuticular.

Asci amphigenous, cylindric, rounded or truncate at the apex;
stalk cells broad, flat, ascospores eight, round, ovate, or elliptic,
frequently budding in the ascus. (Fig. 27, C.)

Dimensions: Of asci, 20-40[jl X 10-13[jl; of stalk cells, 5-7[x X
10-15[jl; of ascospores, 4-5[jl X 4-4.5[jl.

Distribution: Caucasus.

Material examined: Russia: Transcaucasia, Kahetia, 1897, A.

110 The University Science Bulletin

Jaczewski (Herb. Inst. Prot. Plants, Sect. Phytopath. Leningrad.
Part of type).

63. Taphrina amelanchieri Mix
Taphrina amelanchieri Mix, Amer. Jour. Bot. 26:44-48. 1939.

Causing witches' brooms on Amelanchier alnijolia Nutt.

Mycelium subcuticular.

Asci hypophyllous, cylindric to clavate, rounded at apex; stalk
cells variable in size and shape. Ascospores eight, round, ovate, or
elliptic, frequently budding in the ascus. (Fig. 27, D.)

Dimensions: Of asci, 20-34[x X 8-13ij.; of stalk cells, 8-13p. X
6-12[x; of ascospores, 3.5-5.5ix X 3-5[ji..

Distribution: California.

Material examined: California: Siskiyou Co., Yreka, near
Frenchtown, June 24, 1912, E. P. Meinecke (Type) .

64. Taphrina crataegi Sadebeck

Taphrina crataegi Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 8:61-95.
1890.

Exoascus bullatus (Berk.) Fkl, in part. Sadebeck, Jahrb. Hamburg. Wis-
sensch. Anst. 1:93-124. 1884.

Exoascus crataegi Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 10:5-110.
1893.

Causing leaf-curl and deforming shoots (not causing true witches'
brooms) of Crataegus monogyna Jacz., C. oxyacantha L., C. san-
guinea Pall.

Mycelium intercellular.

Asci hypophyllous, cylindric, rounded or truncate at the apex, with
stalk cell ; ascospores eight, round, ovate, or eUiptic, frequently bud-
ding in the ascus. (Fig. 27, E.)

Dimensions: Of asci, 20-36[j. X 8-12ix; of stalk cells, 6-13\). X
6-13[Ji; of ascospores, 4-5. Sf;, X 3.5-4.5[;,.

Distribution: Europe.

Material examined: C. monogyna. Austria: Wienerwald,
Hiihnerberg near Baden, May, 1904, Hohnel (Rehm, Asco. 1612).
Poland: Krakow, Raciborski (Ex. herb. A. Wroblewski).

C. oxyacantha. Germany: Mecklenberg, Kluschenberg near
Stargard, June 5, 1907, P. Sydow (Myc. Ger. 593) ; Saxony, near
Leipzig, May, 1884, Pazschke and Winter (Rab.-Wint. F. Eur.
3057.). Poland: Fredrow, June 20, 1920, A. Wroblewski (Ex. herb.
A. Wroblewski).

C. sanguinea. Russia: Smolensk, 1896, A. Jaczewski (Herb. Inst.
Prot. Plant Sect. Phytopath. Leningrad).

This fungus, included in Taphrina bullata by Fuckel (1873-4),
was separated from that form by Sadebeck (1890).

Mix: Monograph of Genus Taphrina 111

65. Taphrina potentillae (Farl.) Johanson

Taphrina potentillae (Farl.) Johanson, Ofvers, K. Svensk. Vetensk. Akad.
Forhandl. 1885 :29-47. 1886.

Exoascus deformans (Berk.) Fkl. var. potentillae Farlow, N. A. F. 299.
Described by Farlow, Proc. Amer. Acad. Arts and Sci. 18:65-85. 1883.
Taphrina tormentillae Rostrup, Bot. Tidsskr. 14:230-243. 1885.
Magnimella potentillae (Farl.) Sadebeck, Jahrb. Hamburg. Wissensch.
Anst. 10:5-110. 1893.

Causing small, whitish to yellowish thickened areas on leaves and
stems of Geuni montanum L., Potentilla arguta Pursh., P. canaden-
sis L., P. Corsica Lebem., P. flabellifolia Hook., P. geoides L., P.
glandulosa L., P. recta L., P. rwpestris L., P. silvestris Neck., P.
tormentilla Schrk., P. silvestris Neck. X P- procumbens Sibth.,
Potentilla sp.

Mycelium intercellular.

Asci amphigenous, formed beneath epidermis, clavate, rounded or
truncate at the apex, arising from the ends of intercellular hyphae
as though borne on a pedicel; ascospores eight, round, ovate, or
elliptic, usually budding in the ascus (Fig. 27, F.)

Dimensions: Of asci, 17-58[jl X 7-17[jl; of ascospores, 4-5. 5[a X
8-4.51;..

Distribution: Europe, eastern North America.

Material examined: Geum montanum. Corsica: Monte Rotendo,
July 22, 1902, R. Maire (Ex. Herb. R. M.).

Potentilla arguta. Connecticut: West Haven, July, 1890, R.
Thaxter (Rel. Farl. 644). Michigan: East Lansing, June 29, 1943,
E. A. Bessey. Sweden: Uppsala, June 17, 1907, 0. Juel.

P. canadensis. Connecticut: New Haven, June 27, 1928, G. P.
Clinton; 26/(i., June 30, 1928, A. J. M. Massachusetts: Wellesley,
June 8, 1935, D. H. Linder; ibid., June 22, 1935, id. New Jersey:
Newfield, Aug., 1894 (F. Columb. 408). Ohio: Warren Co.,
Foster, May 15, 1937, W. B. Cooke. Tennessee: Elkmont, June 8,
1937, L. R. Hesler. Vermont: Griffith Shelter, Aug. 10, 1936, R. H.
Thompson.

P. Corsica. Corsica: Coscione near Grienza, June 7, 1901, R.
Maire (ex. herb. R. Maire).

P. flabellifolia. California: Mt. Shasta, South Gate Springs,
July 10, 1947, Wm. Bridge Cooke (Herb. W. B. C. 20282).

P. geoides. Sweden: Uppsala, Bot. Gard., June 17, 1907, 0.
Juel; ibid., June 16, 1916, id.

P. glandulosa. California: Mt. Shasta, July 5, 1940, W. B.
Cooke (Herb. W. B. C. 14626).

P. recta. Sweden: Glanshammar, Skafsund, Aug. 21, 1925, T.
Vestergren.

112

The University Science Bulletin

P. rupestris. Germany: Bavaria, Kappellenwiese, near Gerolz-
hofen, Sept. 17, 1915, A. Vill. (Myc. Ger. 1957). Norway: Oslo,
Bot. GarcL, Aug. 24, 1931, I. J0rstad.

P.silvestris. Germany: Brandenburg, Prignitz, Triglitz, Sept. 29,
1898, O. Jaap (F. Sel. Exs. 7). Poland: Krakow, Raciborski (Ex.
herb. A. Wroblewski).

P.procumbens X silvestris. Germany: Schleswig-Holstein, Fohr.
Is., Nieblum, Aug. 1, 1904, O. Jaap (F. Sel. Exs. 7b).

P otentilla sp. [cultiv.). California: Berkeley, March, 1940, H.
N. Hansen.

HOST INDEX TO SPECIES OF TAPHRINA ON
ROSACEAE, EXCLUSIVE OF PRUNUS

Amelanchier alnijolia Nutt.

Taphrina amelanchieii Mix
Crataegus monogyna Jacz.

Taphrina crataegi Sadeb.
Crataegus oxyacantha L.

Taphrina crataegi Sadeb.
Crataegus sanguinea Pall.

Taphrina crataegi Sadeb.
Geum montanum L.

Taphrina potentillae (Farl.) Johans.
Potentilla arguta Pursh.

Taphrina potentillae (Farl.) Johans.
' Potentilla canadensis L.

Taphrina potentillae (Farl.) Johans.
Potentilla Corsica Lebem.

Taphrina potentillae (Farl.) Johans.
Potentilla flabellifolia Hook.

Taphrina potentillae (Farl.) Johans.
Potentilla geoides L.

Taphrina potentillae (Farl.) Johans.
Potentilla glandulosa L.

Taphrina potentillae (Farl.) Johans.
Potentilla recta L.

Taphrina potentillae (Farl.) Johans.
Potentilla rupestris L.

Taphrina potentillae (Farl.) Johans.
Potentilla silvestris Neck.

Taphrina potentillae (Farl.) Johans.
Potentilla tormentilla Schrk.

Taphrina potentillae (Farl.) Johans.
Pyrus amygdalijormis Vill.

Taphrina bullata (Berk.) Tul.
Pyrus betulifolia Bunge

Taphrina bullata (Berk.) Tul.
Pyrus communis L.

Taphrina bidlata (Berk.) Tul.
Pyrus lindleyi Rehd.

Taphrina bidlata (Berk.) Tul.

Taphrina orientalis Mix
Sorbus alnifolia K. Koch.

Taphrina piri Kus.
Sorbus torminalis (L.) Crantz.

Taphrina sorbi (Jacz.) Mix

Mix: Monograph of Genus Taphrina 113

VI. SPECIES ON ROSACEAE

Form on plums: Prunus
66. Taphrina pruni Tulasne

Taphrina pruni Tulasne, Ann. Sci. Nat. 5 Ser. Botanique. 5:122-136. 1866.

Exoascus pruni Fuckel, Jahrb. Nassau Ver. f. Naturk 23 and 24:1-459.

1869 and 1870.

T. rostrupiana (Sadeb.) Giesenhagen, Flora 81:267-361. 1895.

E. rostrupianus Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 10:5-110. 1893.

T. insititiae (Sadeb.) Johanson, Ofvers K. Svensk. Vetensk. Akad. For-

handl. 1885 :29-47. 1886.

E. insititiae Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 1 :93-124. 1884.

T. pruni Tul. var. divaricata Jaczewski, Pocket Key for the determination

of fungi, Part I. Exoascales, Leningrad. 1926.

Causing deformed fruits ("bladder-plums," "plum pockets") and
thickened malformed twigs often bearing curled leaves on Prunus
cerasifera Ehrh. var. divaricata Bailey, P. domestica L., P. insititia
L., P. spinosa L., P. ussuriensis Koval. and Kost.

Mycelium intercellular.

Asci cylindric-clavate, rounded or truncate at the apex; stalk
cell present, sometimes nearly half the length of the ascus. Asco-
spores eight, round, ovate, or elliptic, commonly budding in the
ascus. (Fig. 28, A-F.)

Dimensions: Of asci, 17-53[jl X 5-17pL; of stalk cells, 5-27ix X
4-13[jl; of ascospores, 4-7[jl X 3-6[x.

Distribution: Europe. Japan. (North America?)

Material examined: Prunus cerasifera var. divaricata. Russia:
Caucasus, Suchum, April 27, 1914, W. Siemaszko; Transcaspia,
Kopet-Dagh, Han Ailaion, May 30, 1924, Czerniakowska (det.
Jaczewski).

P. domestica. New Jersey: (slide in Atkinson collection). New
York: Geneva, July 9, 1935, F. C. Stewart. Manitoba: Winnipeg,
July 1, 1935, G. R. Bisby. Japan: Iwate, Tamayama, May 29,
1908, G. Yamada (Herb. Morioka Imp. Coll. Agric. and For.).
Sweden: Bohuslan, Ljings, Korskviken, July 15, 1888, A. G.
Ehasson; Skane, Bunkeflo, June 10, 1884, Eriksson (on shoots),
Smaland, Urasa, Skye, June 3, 1883, Johanson (on shoots) ; Soder-
manland, Allhelgona, Bullersta, June, 1894, G. V. Schotte.

P. insititia {di^ Taphrina insititiae) . Denmark: Sjaelland, Kalle-
have. May 14, 1913, J. Lind (Micr. Rar. Sel. 1744). Germany:
Brandenburg, Triglitz in Prignitz, May 22, 1899, 0. Jaap (F. Sel.
Exs. 51). Hungary: Leutschovia, Locse, June, 1891, V. Greschik
(F. Leutschov.).
8—3659

114

The University Science Bulletin

Fig. 28. Asci (X900) of Taphrina pruni. A, on Prunus domcstica;
B, P. cerasijera var. divaricaia; C, P. ussuriensis ; D, P. domesticn, twigs
(as T. insititiae) ; E, P. spinosa, twigs; F, P. spinosa, fruits (E, F, as T.
rostrujnana) .

Mix: Monograph of Genus Taphrina 115

P. spinosa (as T. rostrupiana) . Dalmatia: Cattara, April, 1914,
0. Jaap (Ex. herb. Sydow, on both fruits and shoots). Italy:
Florence, Vallombrosa, spring 1898, F. Cavara (Myc. Ital. 537),
Poland: Pulawy, May, 1923, W. Konopacka (Herb. Inst. Phyto-
path. Schol. Sup. Varsaviensis). Russia: Nowotscherkassk, May
2, 1911, 0. Trebu (shoots). Sweden: Skane, Kullen, Arild, July
26, 1923, 0. Juel (fruits and shoots) .

P. ussuriensis. Russia: Nicolsk-Ussurisk, July, 1912, N. Naumov
(Herb. Inst. Prot. Plant. Sect. Phytopath. Leningrad).

A portion of a specimen received from R. Maire, labelled:
"Taphrina insititiae, Graecia, in Pruno pseudo-armeniaca, in planitie
Kardianos Kambros monelato-vorno inter Lidiki et Amphissam.
Balais de sorceres effomiat. 14/8 R. Maire" (Miss. Bot. Orient.
1906. 268) was insufficient to allow determination of any fungus
present.

Reduction of these various species to synonymy is based on the
belief (borne out by observation of several other species of Taphrina)
that morphologically similar fungi on the fruits and leaves of the
same host must be the same. Further there is no clear morphological
distinction between different host-forms. The dimensions of the
various fungi, considered as separate species, are as follows:

Asci Stalk cells

T. -pruni on fruits of P. domestica

23-53txX 5-% 11-27;;. X 5-%

T. pruni on fruits of P. cerasifera var. divaricatn

26-53pL X 10-17[x 8-23;jL X 8-12;^

T. insititiae on shoots of P. domestica

17-33|x X 5-10;x 5-10;x X 4-13pL

T. insititiae on shoots of P. insititia

17-33|xX 5-10[/, 5-%X4-%

T. rostrupiana on fruits of P. spinosa

26-53;jlX' 8-% 8-20;x X 5-13;jL

T. rostrupiana on shoots of P. spinosa

23-33[jL X 7- SpL 7-12;;. X 6- Spi,

Taphrina insititiae has shorter asci and shorter, and more nearly
cuboidal stalk cells (often wider than high) than T. pruni. This is
the same sort of difference observed in other pruniculous species be-
tween the asci formed on fruits and those on leaves. The same dif-
ference is to be observed in Taphrina rostrupiana, which has been
considered to be the same fungus whether occurring on fruits or
leaves. There is fairly good agreement in ascus-size between the
three fungi under consideration when occurring on fruits.

The frequently noted occurrence of one of these fungi on fruits

116 The University Science Bulletin

only or on shoots only can be explained by the fact that conditions
necessary to infection of both fruits and shoots may not occur every
season. Magnus (1894) finding plum pockets and malformed shoots
on the same tree of Prunus domestica, and finding no distinction be-
tween the asci from both sources suggested that the same fungus
{T. pruni) was present on both fruits and twigs.

67. Taphrina communis (Sadeb.) Giesenhagen

Taphrina communis (Sadeb.) Giesenhagen, Flora 81:267-361. 1895.

Exoascus communis Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 10 :5-110.

1893.

T. longpipes (Atk.) Giesenhagen, I.e.

E. longpipes Atkinson, Bull. Torr. Bot. Club. 21 :372-380. 1894.

T. decipiens (Atk.) Giesenhagen, I.e.

E. decipiens Atkinson, I.e.

T. rhizipes (Atk.) Giesenhagen, I.e.

E. rhizipes Atkinson, I.e.

E. decipiens Atk. var. superficialis Atkinson, I.e.

T. mirabilis (Atk.) Giesenhagen, I.e.

E. mirabilis Atkinson, I.e.

E. mirabilis Atk. var. tortilis Atkinson, I.e.

Causing deformation of fruits ("bladder plums," "plum pockets")
and of twigs, sometimes with curling of leaves (not witches' brooms)
of Prunus americana Marsh., P. angustifolia Marsh., P. hortulana
Bailey, P. hortulana Bailey var. mineri Bailey, P. lanaia (Sudw.)
Mack, and Bush, P. maritima Marsh., P. rnunsoniana Wight and
Hedr., P. nigra Ait., P. salicina Lindl., P. unibellata Ell.

Mycelium intercellular.

Asci cylindric-clavate, usually rounded at the apex, stalk cell
narrower than the ascus, of variable length, ascospores eight, round,
ovate, or elliptic, frequently budding in the ascus.

Dimensions: Of asci, 27-83[x X 5-13\l; of stalk cells, 6-56[jl X
3-12[jl; of ascospores, 4-7[jl X 3.5-5.5[ji..

Frequently longer asci occur, projecting above the general surface
of the hymenium; these are approximately twice the length of
neighboring asci. Such long asci (which are normal in all other
respects) may reach a length of 116[x. (Fig. 29, A-F, Fig. 30, A-E.)

Distribution: eastern and middle North America.

Material examined: Prunus americana. Arkansas: Conway,
May 1, 1939, T. Jones; Fly Gap (near Cass), May 12, 1940, V. H.
Young, Mountainburg, May 24, 1941, A. J. M. Iowa: (slides, At-
kinson Collection, one on variety Cheney) . Kansas: Abilene, May
22, 1926, H. W. King; Baldwin, May, 1926, A. J. M.; ibid., numerous
collections 1925-1939, id.; Belpre, June 9, 1924, R. P. White; Law-
rence, May 23, 1939, R. Sailer; ibid., K. U. Campus, May, 1942,

Mix: Monograph of Genus Taphrina

117

Fig 29. Asci (X900) of Taphrina communis. A, on Prunus ameri-
rana (fmits) ; B, P. umbellata; C, P. americana (twigs) ; D, P. maritima
(Iruits) ; Jb., P. munsomana; F, P. nigra.

118 The University Science Bulletin

A. J. M.; Rockport, June, 1894, E. Bartholomew (N. A. F. 298);
Rooks Co., May 24, 1901, E. Bartholomew (F. Coliimb. 1533).
Massachusetts: Morden, June 27, 1929, I. L. Conners. Michi-
gan: Alma, June 2, 1894 (slide in Atkinson Collection) ; Forest
Lake, June 18, 1935, 0. J. Eide. New York: Ithaca, Ellis Hollow
Road, June 10, 1940, A. J. M.; ibid., June 22, 1940, id. (fruits and
shoots) ; locality not given (slide in Atkinson Collection, on Bald-
win variety).

As T. decipiens. Kansas: Baldwin, May, 1927, A. J. M.; Stock-
ton, June 1, 1903, E. Bartholomew (F. Columb. 1927). New York:
Aetna, July 6, 1894 (slide in Atkinson Collection). North Caro-
lina: Raleigh, May, 1936, R. F. Poole. Saskatchewan: Indian
Head, Aug. 5, 1926, P. M. Simmonds (Herb. Central Expt. Farms.
149).

As T. decipiens var. superficialis. New York: Aetna, July 6,
1894 (slide in Atkinson Collection).

As T. longpipes. Mississippi: Grand Rapids, 1898, E. J. Coler
(Farl. Herb.). New York: Danby (slide in Atkinson Collection,
from type material) .

P. angustifolia, as T. mirabilis. Alabama: April 28, 1892 (slides
in Atkinson Collection) ; Auburn, April-May, 1890-1892, G. F. At-
kinson (Econ. F. 729). Arkansas: Fayetteville, May, 1940, V. H.
Young; Hamburg, May, 1939, J. C. Dunegan (fruits and shoots).
Florida: Gainesville, March 26, 1936, G. F. Weber; ibid., April 9,
1941, id. Georgia: Athens, May 14, 1936, J. H. Miller. Iowa:
Chicasa Co. (slide in Atkinson Collection). Kansas: 1924, R. P.
White; Lawrence, June, 1923, A. J. M.; ibid., several coll. subse-
quent years, id. Mississippi: Morse, May 21, 1936, L. E. Miles.
Tennessee: Elkmont, June 13, 1937, L. R. Hesler. Virginia:
1935, S. A. Wingard.

As T. mirabilis var. tortilis. Alabama: (slide in Atkinson Collec-
tion). Florida: Gainesville, March 26, 1936, G. F. Weber. Kan-
sas: 1924, R. P. White.

P. hortulana, as T. mirabilis. Arkansas: Bentonville, June,
1907, W. A. Scott (Herb. Cornell Univ. 3502). Iowa: Cedar Rapids
(slide in Atkinson Collection). Wisconsin: Blue River, June 9,
1932, J. J. Davis (Farl. Herb.).

P. hortulana var. mineri. Iowa: 1894, A. Noe (slide in Atkinson
collection).

P. lanata. Arkansas: Conway, May 1, 1939, T. Jones.

P. maritima. Massachusetts: (slide in Atkinson Collection,

Mix: Monograph of Genus Taphrina

119

Fig. 30. Asci (X 900) of Taphrina communis. A, on Prunus maritima
(twigs); B, P. am,ericana {T. rhizipes) ; C, P. americana {T. longipes) ;
D, P. angv^tijolia {T. mirabilis) ; E, P. angustifolia (T. mirabilis var.
tortilis) .

120 The University Science Bulletin

fruits) ; Barnstable, July 13, 1939, 0. C. Boyd (fruits) ; Cape Cod,
June 13, 1939, W. H. Thies and B. Tomlinson (fruits and shoots) ;
Falmouth, June 22, 1940, R. H. Thompson (fruits and shoots) ; West
Falmouth, June 19, 1911, A. B. Seymour (fruits).

P. munsoniana. (Cornell Univ. Herb. 4849, preserved in for-
malin).

P. nigra. Michigan: St. Johns, June 20, 1938, E. A. Bessey.
New York: Ithaca, Highland Road, June 7, 1940, A. J. Eames;
Kline Road, near cemetery, June 7, 1940, A. J. M.; ibid., June 8,
1940, id.; Loon Lake, July, 1940, J. R. Stickney. Ontario: east of
Wilcox Lake, June 14, 1936, R. F. Cain (Herb. Univ. Toronto,
19047). Quebec: Aylmer, June 24, 1937, R. K. Eden (Herb. Cen-
tral Expt. Farms 4614) .

P. salicina. Alabama: Auburn, May 1, 1892, G. F. Atkinson
(apparently type of T. rhizipes, both fruit and shoot affected) ;
ibid., May 8, 1892, id. (three slides in Atkinson Collection). Con-
necticut: Cheshire, June, 1891, R. Thaxter (Herb. R. Thaxter
1497).

P. umbellata. Georgia: Stone Mountain, April 26, 1925, J. H.
Miller (Herb. Univ. Georgia 2475).

As far as Sadebeck (1893) distinguished between Taphrina com-
munis and T. pruni the former had slightly smaller asci and a longer
and narrower stalk cell. Sadebeck's measurements are:

T. pruni, asci 40-55[ji, X 8-15iJi,,stalk cells 10-16[x X 8[x.

T. communis, asci 30-40ij. X 8[x, stalk cells 15-20[j, X 3-5[j..

Actually T. communis has longer and slightly more slender asci
and stalk cells, the dimensions of the two species being as follows:

T. pruni, asci 17-53pi, X 7-17[ji., stalk cells 5-27[a X 4-13[ji,.

T. communis, asci 27-83 (116) jjl X 5-13tJL, stalk cells 7-56[;, X
4-IO1;..

The chief distinguishing feature of T. communis is the occasional
occurrence (mentioned above) of very long asci (as long as 116ijl),
This was observed by Robinson (1887) who spoke of it as "di-
morphism," describing long, slender, usually eight-spored asci,
43-60[A X 5-7[jl; and shorter, thicker, usually many-spored asci,
27-35[JL X 9-12[ji,. Robinson also states that he found the asci of his
"2\ pruni" more slender than those described by Sadebeck. Al-
though the host is given as P. domestica it is apparent that Robinson
was dealing with Taphrina communis. If his host-determination
and those of others who have reported plum pockets on P. domestica
in North America, are correct it means that P. domestica is attacked

Mix: Monograph of Genus Taphrina 121

by T. communis. Apparently it is not highly susceptible. "Domes-
tica" plums are chiefly cultivated in the eastern United States.
Plum pockets on cultivated plums have been less commonly re-
ported from that area than from the Mississippi Valley where
''Americana" varieties are grown. Assuming that Taphrina pruni
and T. communis are separate species it may be doubted that the
former occurs in North America. Plum pockets found on "Domes-
tica" plums in this country should be ascribed to Taphrina com-
munis.

The long asci are quite as common in the form described by At-
kinson as Exoascus mirabilis as in Taphrina communis (in the nar-
rower sense). Indeed the former fungus cannot in any way be dis-
tinguished from T. communis. It deforms shoots of Prunus angus-
tifolia and only rarely attacks fruits. This may be due to a less
degree of susceptibility of the fruits, or to rarity of conditions nec-
essary for fruit infection. In eastern Kansas, Prmvus angustifoUa
seldom bears fruit.

That the fungus on shoots and fruits of P. angustifoUa are one
and the same became clear from a study of the material collected
by J. C. Dunegan at Hamburg, Arkansas. The asci were similar on
both host organs, those from fruits measuring 33-50[ji, X 7-10[;, with
stalk cells 13-23pt. X 5-8pi,, while asci from shoots were 30-46pi, X
7-lOpt,, stalk cells 13-20[x X S-Spt,. (As in other species asci on shoots
are slightly smaller than those on fruits.)

The case for synonymy of Taphrina communis and T. decipiens
has been well presented by Ray (1939) who calls attention to the
fact that asci borne on the leaves of infected shoots are somewhat
smaller than those found on fruits. Shoot infection is, at least in
some localities, less common than fruit infection. In the periods
1926-1929 and 1931-1939 a stand of Prunus americana at Baldwin,
Kansas, was visited annually. Plum pockets occurred every season
but deformed shoots in 1927 only.'^ It is difficult to believe that this
single occurrence of malformed shoots in this isolated plum thicket
was due to a separate fungus. On the other hand, the idea that con-
ditions favoring shoot infection by the fungus {T. communis)
known to be present, may occur but rarely is entirely credible. In the
more humid climate of the eastern United States (Ithaca, New
York, for example) shoot infection seems to be common.

7. The plum thicket in question was not visited for several years after 1939, but in 1947,
an unusually rainy spring, shoot infections (as well as pockets) were observed to be abundant
in all trees. Deformed fruits and shoots were also observed near Lecompton, Kansas, on
May 15, 1947, by R. L. McGregor.

122 The University Science Bulletin

Atkinson (1894) distinguished two other pocket-forming species
from Taphrina communis, namely: Exoascus longipes, and E.
rhizipes. The first of these was characterized by an unusually long
stalk cell. Asci of T. communis with stalk cells as long (SSfji) as
those described by Atkinson for E. longipes, are fairly common and
have been found in many collections. Because of the variability of
Taphrina communis from fruit to fruit of the same collection it
would be impossible to sort out the various specimens studied into
T. communis and "T. longipes."

Atkinson's Exoascus rhizipes, occurring on Japan plum, was char-
acterized by rhizoidal extensions of the lower end of the stalk cell.
Such rhizoidal processes occasionally occur in Taphrina communis
on various hosts and are no more abundant in Atkinson's type ma-
terial of E. rhizipes than in many other collections.

68. Taphrina pruni-subcordatae (Zeller) Mix

Taphrina ■pruni-s.uhcordatae (Zeller) Mix, Univ. of Kansas Sci. Bull. 24:10:

151-176. 1936.

Exoascus pruni-subcordatae Zeller, Mycologia 19:130-143. 1927.

Causing deformed fruits (plum pockets) and thickened, mal-
formed shoots on Prunus subcordata Benth.

Mijcelium intercellular.

Asci clavate, rounded or truncate at the apex, provided with a
stalk cell, ascospores eight, round, ovate, or elliptic, often budding in
the ascus (Fig. 31, A, B).

Dimensions: Of asci, 33-73[x X '7-12[j(.; of stalk cells, 7-40tx X
5-12pL; of ascospores, 4-7[j. X 3.5-6[jl.

Distribution: Rocky-Mountain and Pacific-Coast areas, North
America.

Material examined: California: Cordelia, April 22, 1936, H.
Earl Thomas (fruits) ; Elk Creek, May 15, 1938, id. (fruits) ; ihid.
(a few miles distant). May 15, 1938, id. (shoots); Mt. Shasta,
Sept. 6, 1940, W. B. Cooke (Herb. W. B. C. 14724, shoots) ; Tay-
lorsville, April 23, 1915, J. S. Boyce (Herb. J. S. B. 285, shoots).
Colorado: Boulder, 1942, A. J. M. (fruits and shoots) ; Fort Col-
lins, 1935, L. W. Durrell (fruits).

This species is closely related to Taphrina communis. Its asci
are usually somewhat longer and wider and the stalk cells a little
shorter than in that species. The exceptionally long occasional asci
characteristic of T. communis are absent.

Mix: Monograph of Genus Taphrina

123

Fig. 31. Asci (X900) of A, B, T. pruni-subcordatae (A, on fruits;
B, on twigs); C, D, T. mume; E, T. deformans on Prunus communis;
F, G, T. deformans on P. persica; H, T. mexicana; I, T. armeniacae.

124 The University Science Bulletin

Forms on Apricots

69. Taphrina mume Nishida

Taphrina mume Nishida. Miyabe Festschrift Tokyo, 1911.

Taphrina deformans (Berk.) Fkl. var. armeniaca Ikeno, Flora 92:1-31. 1903.

Causing leaf-curl and hypertrophy of twigs of Pninus armeniaca
L. var. ansu Maxim., P. mandshurica Koehne, and P. viume Sieb.
and Zucc.

Mycelium intercellular.

Asci amphigenous or hypophyllous, cylindric, rounded or truncate
at the apex, with a broad, short stalk cell; ascospores eight, round,
ovate, or elliptic. (Fig. 31, C, D.)

Dimensions: Of asci, 20-33[ji. X 8-13[;i. (23-52[x X 8-15[x according
to Nishida); of stalk cells, 7-15[jt. X 5-15pi.; of ascospores, 4-6[;, X

Distribution: China, Japan.

Material examined: P. armeniaca. Japan: Fukui, Togo-mura,
May 18, 1922, T. Toyoda (Herb. Morioka Imp. Coll. Agric. and
For.)

P. mume. Japan: Hiroshima, May 7, 1930, S. Kusano; Iwate,
Morioka, June 23, 1927, K. Togashi (Herb. Morioka Imp. Coll.
Agric. and For.) ; ibid., June 17, 1930, G. Yamada id.

This species was considered by Ikeno (1903) and also by Tai
(1937) to be a variety of Taphrina deformans. It obviously is
closely related to Taphrina deformans, and if Nishida's measure-
ments are correct, is not morphologically distinguishable from that
fungus. Though it is probably identical with T. deformans, it is
retained as a separate species pending further study.

The occurrence of Taphrina mume on Prunus mandshurica is
reported by Tai. No specimen has been seen by the writer.

70. Taphrina arm^eniacae Georgescu and Badea

Taphrina armeniacae Georgescu and Badea, Analele Institului de Cercetari
si Experimentatie Forestura 3:162-167. 1938.

Causing witches' brooms on Prunus armeniaca L.

Mycelium intercellular.

Asci amphigenous, cylindric-clavate, rounded or truncate at the
apex, a stalk cell usually present but occasionally absent; asco-
spores eight, round, ovate or elliptic.

Dimensions: Of asci, 22-29[jl X 7-10[x; of stalk cells, 7-lOpL X
7-13!Ji; of asci lacking stalk cells, 29-38[x X 8-10[ji,; of ascospores,
4-7[.X4-6i.. (Fig. 31,1.)

Mix: Monograph of Genus Taphrina 125

Distribution: Rumania.

Material examined: Twigs and leaves preserved in alcohol, re-
ceived June, 1947, from Botanical Laboratary, Polytechnic Insti-
tute, Bucharest.

Georgescu and Badea state that this fungus causes a witches'
broom, with negatively geotropic curvature of twigs. The material
studied may well have been from such a witches' broom, but this
could not be detemiined with certainty. The affected leaves are
not curled nor conspicuously thickened.

Georgescu and Badea also describe the asci as hypophyllous and
lacking a stalk cell. In the material studied asci were regularly
amphigenous, and all but a few asci possessed well defined stalk
cells. Only by careful search could an occasional ascus lacking a
stalk cell be found.

71. Taphrina deformans (Berk.) Tulasne

Taphrina deformans (Berk.) Tulasne, Ann. Sci. Nat. 5 Ser. Bot. 5:122-136.

1866.

Exoa&cus deformans (Berk.) Fuckel, Jahrb. Nassau. Ver. Naturk. 23 and

24:1-459. 1869 and 1870.

Ascomyces deformans Berkeley, Outl. Brit. Fungi. 1860.

Taphrina amygdali (Jacz.) Mix, Univ. Kansas Sci. Bull. 24:10:151-176.

1936.

Exoascus am.ygdali: Jaczewski, Pocket Key for Determination of Fungi.

Part I. Exoascales. Leningrad. 1926.

Causing leaf curl and malformation of twigs of Prunus communis
Arcang. var. amara Schneid., var. dulcis Schneid., and P. persica L.

Mycelium intercellular.

Asci epiphyllous, occasionally amphigenous, cylindric-clavate,
rounded or truncate at the apex, provided with a stalk cell. As-
cospores eight, round, ovate, or elliptic, frequently budding in the
ascus. (Fig. 31, E, F, G.)

Dimensions: Of asci, 17-56[jl X 7-15ijl; of stalk cells, 6-20[ji, X
5-15pL; of ascospores, d-7\i. X 3-7ijl.

Distribution: world wide, apparently coincident with that of its
hosts.

Material examined: P. persica. California: Berkeley, May 17,

1935, Ruth F. Allen ("ornamental peach") ; Palo Alto, April 21,
1939, R. H. Thompson (P. persica var. plena) ; ibid., Stanford Univ.
orchard. May 23, 1895, S. H. Burnham (nectarine). Illinois:
Fayette Co., May 14, 1936, G. H. Boewe; Johnson Co., May 11,

1936, id. Kansas: Lawrence, numerous collections 1924-1939, A. J.
M. Michigan: Ann Arbor, May 28, 1933, E. B. Mains (Herb.
Univ. Michigan). North Carolina: May, 1936, R. F. Poole. Vir-

126 The University Science Bulletin

ginia: Norfolk, May 6, 1936, H. T. Cook. Ontario: London, June,
1892, J. Dearaess (F. Columb. 139). China: Kwangsi, Long Ping
(Ling Yuin Hsein), April 9, 1933, S. Y. Cheo (Herb. Univ. Nan-
king). England: 1938 (material in alcohol from Royal Coll. of
Sci., London). Japan: Iwate, Kadoma, June 15, 1931, K. Togashi
(Herb. Morioka Imp. Coll. Agric. and For.) ; Morioka, College
Orchard, July 1, 1931; K. Togashi id. Poland: Pulawy, 1923,
W. Konopacka (Herb. Inst. Phytopath. Scol. Sup. Agric. Varsa-
viensis).

P. communis. England: specimens in alcohol received from
Prof. W. Brown, Royal Coll. of Sci., London. Russia: Caucasus,
Derbent, June 10, 1930, A. S. Letov and L. S. Gutner (Herb. Inst.
Prot. Plant. Sect. Phytopath. Leningrad). Scotland: Aberdeen,
June 19, 1939, Prof. Matthews; St. Andrews, July, 1938, J. A. Mac-
donald (in alcohol), ibid., June 20, 1939, id.

The form on Prunus communis, whose occurrence was first re-
corded by Rathay (1878), has usually been considered identical
with Taphrina deformans. Campbell (1925) called it a variety of
T. deformans, stating that its asci were without stalk cells. In this
he was in error. The material received from J. A. Macdonald was
collected at St. Andrews from the tree from which Campbell ob-
tained his specimens. This material showed typical asci with stalk
cells.

Jaczewski (1. c.) in describing Exoascus amygdali stated that it
was different from Campbell's fungus (his statement being founded
on Campbell's mistaken description) . Jaczewski also stated that
the asci were regularly hypophyllous. Jaczewski's type material
v/as not obtainable but material that he had examined was re-
ceived from K. Naumov. In these specimens asci were mostly
epiphyllous but occasionally amphigenous.

Cultures were secured in 1939 from the specimens collected by
Matthews at Aberdeen, and by Macdonald at St. Andrews. In both
cases the material was received in London the day following its
collection and isolations were made in the laboratory of Prof. W.
Brown of the Royal College of Science.

Using these cultures inoculations of peach were made at Law-
rence in the spring of 1947. The trees chosen had been planted in
a campus nursery three years previously and had never shown any
curl. They were sprayed with 1-50 formalin a few weeks before
inoculation. On April 2 during a light rain, inoculum from agar
cultures was smeared on opening buds. (Green tips were protrud-

Mix: Monograph of Genus Taphrina 127

ing about an eighth of an inch from these buds and natural infec-
tions of unsprayed trees in the neighborhood had undoubtedly oc-
curred during a rainy period a few days earlier.) Several lower
branches of one tree were inoculated with cultures from almond,
those of another with cultures from peach. Severe curl resulted
in both trees. It appeared later than natural curl on unsprayed
trees and asci were not quite mature on May 20, though asci from
natural curl had matured as early as May 10. Sprayed but un-
inoculated branches in the upper parts of these trees remained free
from curl.

This indicates that the fungus on Prunus communis is not bio-
logically distinct from the form on P. persica and that the former
should not be distinguished from Taphrina deformans either as a
separate species or as a variety.

72. Taphrina mexicana H. and P. Sydow
Taphrina mexicana H. and P. Sydow, Ann. Mycol. 18:154-160. 1920.

Causing small witches' brooms on Prunus microphylla (HBK)
Gray.

Mycelium intercellular.

Asci hypophyllous, clavate, rounded or truncate at the apex,
stalk cell usually narrower than the ascus; ascospores eight, round,
ovate, or elliptic, small, often budding in the ascus. (Fig. 31, H.)

Dimensions: Of asci, 22-30t/, X 7-10[x; of stalk cells, S-l'o\i X
6-8[j,; of ascospores, 3.5-4[jl X 2.5-3. Sfx.

Distribution: Mexico.

Material examined: Mexico, 1913, C. Reiche (Bot. Mus. Berlin.
Type).

Forms on Cherries: Microcerasus

73. Taphrina cerasi-microcarpae (Kuschke) Laubert

Taphrina cerasi-microcarpne (Kuschke) Laubert, in S'orauer, Handb.

Pflanzenkr. 2:457-499. 1928.

Exoascus cerasi-microcarpae Kuschke, Monit. Jard. Bot. Tiflis 31 :23-27.

1913.

Deforming fruits (causing "pockets") of Prunus microcarpa C. A.
Mey., P. tomentosa Thunb. Perhaps also deforming twigs of P.
humilis Bunge.

Mycelium intercellular.

Asci cylindric-clavate, rounded at the apex, provided with a stalk
cell; ascospores eight, round, ovate, or elliptic, frequently budding
in the ascus. (Fig. 32, A.)

Dimensions: Of asci (P. microcarpa) , 35-50tx X 10-13[jl (Kuschke),

128 The University Science Bulletin

(P. tomentosa) 26-46pL X 7-lOpi. (Mix) ; of stalk cells (P. micro-
carpa) "short" (Kuschke), 5-l[i. long (Jaczewski 1926), (P. tomen-
tosa) 8-13[i, X 7-ll[ji. (Mix) ; of ascospores, 5-7.5[ji. X 5-6.25[i.

Distribution: Caucasus, Japan, China.

Material examined: P. tomentosa. Japan: Ishikari, Hokkaido,
June 30, 1902, G. Yamada (Herb. Morioka Imp. Coll. Agric. and
For.).

Efforts to obtain a specimen of -the fungus on Prunus microcarpa
from Tiflis or from Leningrad were unsuccessful. It is not known
whether a type specimen exists. Some specimens must have been
available to Jaczewski (1926). Since the fungus has not been seen
the description given above is taken from Kuschke (1. c.) and from
Jaczewski (1926).

The fungus on P. tomentosa (called by its collector T. pruni) is
placed here because of its similarity (except for somewhat narrower
asci and longer stalk cells) and because the hosts are closely re-
lated. For the latter reason the fungus on P. humilis reported by
Tai (1937) and called by him Taphrina truncicola is also tenta-
tively assigned to T. cerasi-microcarpae.

74. Taphrina fiavorubra Ray
Taphrina fiavorubra Ray, Mycologia 31:56-75. 1939.

Causing smallish, elongated, pointed "plum pockets," and thick-
ened deformed shoots on P. besseyi Bailey, P. pumila L., P. pumila
L. var. susquehanae Jaeg.

Mycelium intercellular.

Asci cylindric-clavate, rounded at the apex, provided with a
stalk cell; ascospores eight, round, ovate, or elliptic, frequently bud-
ding in the ascus. (Fig. 32, B, C).

Dimensions: Of asci, 20-53iJi, X 6-12[jl; of stalk cells, 8-26[jl X
4-10[ji,; of ascospores, 3-8[ji. X 2-5[x.

Distribution: North America.

Material examined: P. besseyi. Kansas: Stockton, May 20, 1889,
E. Bartholomew (N. A. F. 288). Nebraska: Eagle, May 13, 1941,
R. W. Goss. Oregon: Corvallis, May 11, 1939, S. M. Zeller. Prince
Edward Island: Little York, Coolhead Road, Mrs. W. J. Mc-
Donald (Herb. Central Exp. Farms. 4610).

P. pumila. Kansas: Rooks Co., 1893, E. Bartholomew (F. Co-
lumb. 924). Maine: Fort Kent, July 1, 1904, M. L. Fernald
(Maine Flora, Aroostook County). Michigan: Arenas Co., July
20, 1938, E. A. Bessey. South Dakota: Brookings, June, 1893,
Griffiths.

Mix: Monograph of Genus Taphrina

129

Fig. 32. Asci (X 900) of A, Taphrina ccrasi-microcarpae ; B, C, T.
flavorubra ; B, on Prunu^ beaaeyi; C, P. pumila var. susquehanac ; D, E,
T. Iruncicola ; D, on Prunus incisa; E, P. maximowiczii.

9—3659

130 The University Science Bulletin

P. pumila var. susquehanae. New York: Ithaca, South Hill
Marsh, June 9, 1937, W. W. Ray (Herb. Dept. Plant Path. Cornell
Univ. 26425. Fruits. Type) ; ibid., June, 1940, A. J. M.; ibid., June
1946, id. Wisconsin: Millston, June 9, 1914, W. W. Davis; Ne-
cedah, June 9, 1917, id. (Herb. Marquette Univ.).

Ray (1939) described this species on Prunus puinila L. var.
susquehanae (Willd.) Jaeg. The fungi on the closely related P.
pumila L., and P. besseyi Bailey agree closely in morphology with
T. flavorubra, and cause similar host-deformations. It seems proper
to assign them to this species.

Forms on Cherries: Pseudocerasus, Lobopetalum, Eucerasus,
Mahaleb, and Phyllomahaleb

75. Taphrina truncicola Kusano
Taphrina truncicola Kusano, Bot. Mag. Tokyo 19:1-5. 1905.

Causing thickening and malformation of shoots (not witches'
brooms) and deformation of inflorescenses of Prunus incisa Thunb.,
and P. maximowiczii Rupr.

Mycelium intercellular.

Asci cylindric-clavate, rounded at the apex, provided with a stalk
cell. Ascospores eight, round, ovate, or elliptic, frequently budding
in the ascus. (Fig. 32, D, E.)

Dimensions: Of asci, 30-45[;, X 7-13[jl; of stalk cells, 10-23[jl X
5-12[;.; of ascospores, 4.5-6. 5[jt, X 4-5. 5[j,.

Distribution: Japan.

Material examined: P. incisa. Japan: Iwate, Mt. Iwate, June
18, 1908, G. Yamada (Herb. Morioka Imp. Coll. Agric. and For.) ;
ibid., June 4, 1919, id.

P. maximowiczii. Japan: Iwate, Mt. Iwate, June 14, 1919, F.

Sawada (Herb. Morioka Imp. Coll. Agric. and For.) ; Nikko, June,

1900, T. Makino.

76. Taphrina cerasi (Fkl.) Sadebeck

Taphrina cerasi (Fkl.) Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 8:61-
95. 1890.

FJxoascus deformans f. cerasi Fuckel, Jahrb. Nassau. Ver. Naturk. 23- and
24:1-459. 1869 and 1870.

E. cerasi (Fkl.) Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 10:5-110.
1893.

E. wiesneri Rathay, Oesterreich Bot. Zeitschr. 30:225. 1880. (No descrip-
tion.)

T. gilgii Hennings and Lindau, Hedwigia 32:156-157. 1893.
T. minor Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 8:61-95. 1890.
E. minor Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 10:5-110. 1893.
T. pseudocerasi (Shirai) Saccardo, Sylloge Fungorum 14:824.
Taphria pseudocerasi Shirai, Bot. Mag. Tokyo 9:161-164. 1895.
E. pruni-acidae Jaczewski, Pockey Key for the determination of fungi.
Part I. Exoascales. Leningrad. 1926.

Mix: Monograph of Genus Taphrina 131

Causing leaf curl (with only slight thickening) and witches'
brooms on Primus avium L., P. cerasus L., P. cerasus L. var,
frutescens (Neilr.) Schneid. (P. acida K. Koch.), P. fruticosa
Pall., P. pennsylvanica L., P. pseudocerasus Lindl., P. sernilata
Lindl. var. lannesiana Rehd., and var. spontanea Wils., P. yedoensis
Matsum.

Mycelium intercellular, perennial.

Asci hypophyllous, rarely amphigenous, clavate, rounded at the
apex, provided with a stalk cell. Ascospores eight, round, ovate,
or elliptic, often budding in the ascus. (Fig. 33, A-F.)

Dimensions: Of asci, 17-53[x X 5-15[x; of stalk cells, 5-26[x X
4-12[jl; of ascospores, 3.5-9[jl X 3-6^/,.

Distribution: Europe, North America, Australia, New Zealand,
South Africa, Japan.

Material examined: Primus avium. California: Berkeley, June
6, 1935, H. Earl Thomas; Napa, April 21, 1929, id. New York:
Ithaca, Coddington Road, May 27, 1937, W. W. Ray; near R. 0.
T. C. Stables, May 19, 1938, id.; ibid., May 31, 1940, A. J. M.
Oregon: Hood River, May 16, 1936, J. R. Kienholz; Polk Co.,
June 17, 1935, S. M. Zeller. Washington: Western Washington
Exp. Sta. 1935. Czechoslovakia: Prencow, Teplicky, June 20,
1890, A. Kmet (Fungi Schemnitz). England: Gloucestershire,
Woodchester Park, May 13, 1934, E. M. Wakefield (Herb. Hort.
Bot. Reg. Kew). Germany: Saxony, Putzkau, June 11, 1908, A.
Schade (Myc. Genu. 798). Norway: Hordaland, Alsaker in Kin-
sarvik, June 5, 1929, I. J0rstad. Poland: Pulawy, June, 1923, Z.
Czarnocka (Herb. Inst. Phytopath. Sch. Sup. Varsaviensis). Rus-
sia: Lublin, Nowo-Alexandria, June 1, 1910, H. Newodowski.
Sweden: Uppland, Uppsala, June 2, 1895, A. G. Eliasson.

P. cerasus. British Columbia: Vancouver Island, Coutenay,
June 8, 1941, W. S. Jones. Germany: Berlin, Grossbeeren, July,
1893, Lindau and Gilg (Herb. Sydow in Bot. Mus. Stockholm as
"forma Gilgii"). Lithuania: Kaunas, Aliksotas, May 29, 1936,
A. Minkevicius (Herb. Univ. Lithuania). Russia: Kursk, May,
1916, A. Bondarzew (F. Ross. Exs.).

P. cerasus var. jrutescens. Germany: Berlin, Zehlendorf, May
9, 1911, R. Laubert (type of T. pruni-acidae) ; Saxony, Islebia,
June, 1875, J. Kunze (Kunze, F. Sel. Exs. 168 as "£?. deformans
f. cerasi-acidae , forma nova").

P. fruticosa (as T. minor). Czechoslovakia: Kromau, May,
1913, H. Zimmerman (Fl. Boh. and Mor. Exs. II, 1, 19942). Ger-

132

The University Science Bulletin

Fig. 33. Asci (X900) A-F, of Taphrina cerasi. A, on Prunus avium;
B, P. cerasus; C, P. fruticosa {T. minor); D, P. pennsylvanica ; E, P.
pneudoccrasus; F, P. yedoensin ; G, asci of T. flecians.

Mix: Monograph of Genus Taphrina 133

many: Hamburg, Winterhude, June 14, 1896, 0. Jaap (F. Sel.
Exs. 8.). Poland: Skomorochy, 1917, Wroblewski.

P. pennsylvanica. Colorado: Boulder Canyon, El Vado, July
1-2, 1941, A. J. M. Maine: Pembroke, June 9, 1934, J. R. Hans-
brough (Herb. For. Path. 81694). Minnesota: Bear River, Big
Bear Lake, June 29, 1947, A. J. M. New Hampshire: Cherry
Mountains, near Twin Mountain, June 13, 1923, J. S. Boyce (Herb.
J. S. B. 1849) ; Columbia, June 19, 1936, H. G. Eno (Herb. For.
Path. 69964); New York: Cranberry Lake, June 12, 1926, P.
Spaulding (Herb. For. Path. 16835) ; Ithaca, Connecticut Hill, June
7, 1940, A. J. M.; near Ringwood, May 28, 1937, W. W. Ray; Wat-
kins Glen, June 6, 1940, A. J. M. North Carolina: Nantahala
Gorge, May 29, 1941, A. J. M. Manitoba: Winnipeg, June 25,
1935, G. R. Bisby; east of Beausejour, June 23, 1947, A. J. M.
Ontario: Tillsonburg, French Farm, May 19, 1938, P. G. Newell.
Quebec: Duchesnay, June 13, 1939, R. Pomerleau.

P. pseudocerasus (as T. pseudocerasi) . Japan: Ishikari, Sapporo,
June, 1902, G. Yamada (Herb. Morioka Imp. Coll. Agric. and For.) ;
Komaba, Coll. of Agric, May, 1900, S. Kusano; Tokyo, May, 1899,
Shirai (Herb. Sydow in Bot. Mus. Stockholm).

P. serndata var. spontanea (as T. pseudocerasi) . Japan: Iwate,
Tsunagi, May 17, 1908, G. Yamada (Herb. Morioka Imp. Coll.
Agric. and For.) ; Morioka, Kuroishino, May 25, 1904, G. Yamada
[id.).

P. yedoensis. Japan: Iwate, Morioka, May 6, 1931, K. Togashi
(Herb. Morioka Imp. Coll. Agric. and For.) ; ibid., May 19, 1934,
id. id.

The forms here included are alike in size and shape of asci and
in size of spores. Spores are seldom of diagnostic value in species
of Taphrina but the ascospores of T. cerasi (and of the related T.
flavorubra) are exceptionally large.

These fungi occur on closely related host-species, causing on all
of them a characteristic leaf curl and on all hosts but two, perennial
witches' brooms with clustered twigs showing negatively geotropic
curvature.

Taphrina minor on Prunus fruticosa (P. chamaecerasns) was de-
scribed by Sadebeck (1890) as causing leaf-curl but no witches'
brooms. The fungus described by Laubert (1912) as "Taphrina
sp." and named by Jaczewski (1926) Exoascus pruni-acidae did
not cause a witches' broom. It occurred on adventitious shoots aris-
ing from roots of Prunus cerasus. The host was called by Laubert

134 The University Science Bulletin

P. acida K. Koch, and he stated that it was not the "Glaskirsche"
F. cerasus var. acida Shrk. (P. cerasus L. var. carproniana L.).
Laubert- suggests that Sadebeck in describing Taphrina minor may
have mistaken a wild form of Primus cerasus for Prunus fruticosa.

Just who was responsible for first ascribing to Taphrina minor
leaf curl of Prunus avium and P. cerasus, distinguishing it from
witches' brooms caused by T. cerasi on the same hosts, has not been
learned, but the practice has become rather common, especially
among English authors. In the Farlow Herbarium a collection of
Taphrina cerasi bears the following in Farlow's handwriting: "Ex-
oascus minor Sad. on cultivated cherry. Deal, England, June, 1899.
No Hexenbesen. Piece sent to Sadebeck." No letter from Sade-
beck later than 1896 could be found among Farlow's correspond-
ence. Sadebeck's opinion on this specimen would have been inter-
esting since he never reported T. minor on Prunus avium or on P.
cerasus.

Leaf curl of cultivated cherry is apparently the first stage in the
establishment of a witches' broom. It may occur in trees showing
no witches' brooms (collection of H. Earl Thomas at Napa, Cali-
fornia, 1929) or in trees showing witches' brooms (collection of W.
W. Ray at Ithaca, N. Y., 1938).

Why Atkinson (1894) ascribed the fungus on Prunus pennsyl-
vanica to Exoascus insititiae is hard to understand. That fungus
occurs on species of Prunus (P. domestica, P. insititia) not closely
related to P. pennsylvanica, and causes twig malformations not at
all resembling witches' brooms.

Taphrina pseudocerasi was never a well distinguished species and
apparently was not recognized by Nishida (1911), who records
Taphrina cerasi on Prunus serrulata Lindl. var. spontanea Maxim.,
and on P. subhirtella Miq. (the hosts for T. pseudocerasi).

The morphological similarity of the various host-forms of Taph-
rina cerasi may be seen from the following tabulation.

Taphrina cerasi on:

Asci Stalk cells

Prunus avium 20-53[j(, X 7-15[jl 6-20p(, X 5-10[ji,

Prunus cerasus 20-33^;, X 7- 8|j, 7-13[j[. X 5- 8^

Prunus cerasus var. jrutescens 20-36pi, X 7-10[x 8-17[x X 5- Opi,

Prunus fruticosa 20-43[jl X 7-12jj^ 7-17[ji, X 5-10[ji,

Prunus pennsylvanica 20-46fji, X 7-13[ji. 6-20[ji, X 5-10[ji,

Prunus pseudocerasus 17-33[jl X 7-lOpi, 7-17[jl X 7-10[ji,

Prunus serrulata 17-30[jl X 7-13[i, 8-17[x X 5-10[x,

Prunus yedoensis 20-36^ X 5-10^ 5-12[x X S-lOpi,

Mix: Monograph of Genus Taphrina 135

It will be seen that the asci and stalk cells are largest on Prunus
avium, but more specimens of the fungus on this host were examined.
Also it is evident that Taphrina minor cannot be distinguished from
T. cerasi by its smaller size, or if so distinguished, T. minor should
include the fonn on P. cerasus and also the forms on Japanese
cherries. (Sadebeck gave as length of asci, for T. minor 30-35ix,
and for T. cerasi SO-SOjjl.

The occurrence of Taphrina cerasi on Prunus serrulata var. lanne-
siana is based on the report of Rathbun-Gravatt (1927) who, how-
ever, did not actually determine the fungus from this host, but only
from P. yedoensis.

77. Taphrina flectans Mix
Taphrina flectans Mix, Amer. Jour. Bot. 26:44-48. 1939.

Causing leaf-curl and witches' brooms of Prunus emarginata
Walp.

Mycelium intercellular, perennial.

Asci hypophyllous, clavate, rounded at the apex, provided with
a stalk cell. Ascospores eight, round, ovate, or elliptic, often bud-
ding in the ascus. (Fig. 33, G.)

Dimensions: Of asci, 20-43[jl X 7-12[jt,; of stalk cells, S-lSpi X
5-12[ji,; of ascospores, 4.5-6[ji. X 4-5[jt..

Distribution: Pacific Coast region of North America.

Material examined: California: near Donner Lake, July 13,
1938, H. N. Hansen; Mt. Sanhedrin, Sept. 4, 1939, C. G. Thompson;
Mt. Shasta, July 7, 1939, W. B. Cooke (Herb. W. B. C. 13323) ;
ibid., June 16, 1939, id. (Herb. W. B. C. 13255) ; Plumas Co., Gold
Lake Lodge, July 31, 1942, Lee Bonar. Idaho: Moscow, June 9,
1935, (received from J. Dearness). Oregon: Oregon Caves, Lake
Mountain Trail, Aug. 12, 1929, G. D. Darker.

Much hesitation was felt at the time this fungus was described
as a separate species (Mix, 1939). It is morphologically similar
to Taphrina cerasi, and its ascus-dimensions fall within the size-
range observed in that species. Predominantly the asci of T.
flectans are somewhat shorter and broader. Consistent treatment
might call for the inclusion of this form within Taphrina cerasi or
for the removal of the fungus on Prunus pennsylvanica (and of
perhaps other host-forms) from Taphrina cerasi and erection of a
new species therefrom. A knowledge of biological relationships
within the group of forms occurring on Cerasus would be helpful.

136

The University Science Bulletin

Fig. 34. Asci (X 900) of A, Taphrina jarlowii; B, C, D, T. andina,
C, {T. reichei), D, on fruits; E-H, Taphrina confusa, E, on P. virginiana,
F, P. demissa, G, P. melanocarpa, H, P. alabamends.

Mix: Monogkaph of Genus Taphrina 137

Forms on Cherries: Padus
78. Taphrina farlowii Sadebeck

Taphrina jarloxmi Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 8:61-95.

1890.

Exoascus jarlowii Sadebeck, Jahrb. Hamburg. Wissensch. Anst. 10:5-110.

1893.

Taphrina varia (Atk.) Mix, Univ. Kansas Sci. Bull. 24:10:151-176. 1936.

E. varixis Atkinson, Bull. Torr. Bot. Club 21 :372-380. 1894.

Causing leaf curl, thickened, malformed twigs (not witches'
brooms), and deformed fruits (pockets) on Prunus serotina Ehrh.

Mycelium intercellular.

Asci epiphyllous, or covering surface of fruit, clavate, rounded at
the apex, provided with a stalk cell. Ascospores eight, frequently
budding in the ascus, round, ovate, or elliptic. (Fig. 34, A.)

Dimensions: Of asci on fruits, 13-33pL X 7-13[ji,, stalk cells, 7-13[x
X 7-12[x; asci on leaves, 13-40[x X 7-13[x; stalk cells, 6-13[jl X 5-10[ji;
of ascospores, 4.5-6[x X 3.5-5[jl.

Distribution: eastern North America.

Material examined: Alabama: Auburn, May, 1891 (Econ. Fungi
128, leaves) ; ibid., April 29, 1892, slide No. 93 of Atkinson col-
lection (young stem). Arkansas: Fayetteville, April 28, 1938, J.
C. Dunegan (leaves, twigs, and fruits). Florida: Gainesville, April
28, 1941, G. F. Weber (leaves). Georgia: Athens, April 22, 1936,
J. H. Miller (leaves). North Carolina: Raleigh, May 11, 1936,
R. F. Poole (leaves). Vermont: Peru, Stark Monument, July 27,
1927, P. Spaulding (For. Path. 16177, fruits) ; Smuggler's Notch,
June 22, 1927 (For. Path. 16129, leaves). Germany: Charlotten-
burg, Schlossgarten, Aug. 6, 1891, P. Sydow (Myc. March. 3459,
fruits). Locality not given. May 29, 1892, G. F. Atkinson (slide
in Atkinson collection, twig).

The identity of the fungus on fruits and shoots is readily seen
in the material collected in Arkansas by Dunegan. Not only is the
fungus similar on all host-parts, but obviously all the asci arise
from the same mycelium.

The occurrence of Taphrina farlowii in Germany (Sydow's speci-
men) must mean that the fungus was introduced with its host.

79. Taphrina andina Palm

Taphrina andina Palm, Svensk. Bot. Tidskr. 3:192-195. 1909.

T. reichei Werdermann, Notizbl. Bot. Gart. and Mus. Berlin-Dahlem.

8 :221-222. 1922.

T. atkinsonii Ray, Mycologia 31 :56-75. 1939.

Causing leaf-curl, witches' brooms, and tremendously elongated
(to 6 cm.) fruit-deformations (pockets) on Prunus serotina Ehrh.
var. salicifolia Koehne (P. capuli Cav., P. capoUin Koehne) .

138 The University Science Bulletin

Mycelium intercellular.

Asci amphigenous on leaves, also covering fruits and floral parts,
cylindric, rounded at the apex, provided with a stalk cell which is
variable in length, somewhat narrower than the ascus, and rounded,
or truncate and sometimes broadened below. Ascospores eight,
round, ovate, or elliptic, often budding in the ascus. (Fig. 34, B,
C, D.)

Dimensions: Of asci, 23-40[jl X T-lSpt,; of stalk cells, 7-26[;i, X
5-12[ji,; of ascospores, 4-5.5[ji. X 3.5-4.5iji,.

Distribution: Ecuador, Mexico.

Material examined: Ecuador: Valle de Chillo, Nov. 13, 1924,
F. L. Stevens (Fungi of Ecuador 288, leaves). Mexico: near City
of Mexico, 1920-1921, C. Reiche (Bot. Mus. Berlin, type of T.
reichei, leaves) ; Valley of Mexico, June 5, 1896, C. C. Pringle (Farl.
Herb., witches' broom) ; Durango, San Ramon, April 21-May 8,
1906, E. Palmer (Plants of Mexico 173, fruits and floral parts). No
locality or date (slide in Atkinson collection).

The type specimen of Taphrina andina has apparently been lost.
According to Patouillard and Lagerheim (1895) the fungus was col-
lected by Lagerheim who, in giving it to Palm, stated that he had
not seen a witches' broom on the host. Palm (1909) makes the lack
of a witches' broom his chief reason for distinguishing the fungus
from Taphrina cerasi. In 1939 no specimen of T. andina could be
found among Lagerheim's collections at Stockholm's Hogskola. The
collection by Stevens corresponds well with Palm's description of
T. andina.

There is no reason for considering the fungus causing leaf curl
and witches' brooms as different from that deforming fruits and
floral parts. The asci on fruits are longer and narrower (26-40[j, X
1-10^) than on leaves (23-36t;. X 8-14[a) . Stalk cells from fruits
are also longer and narrower (10-26pi, X 5-9[ji) than those from leaves
(7-23[i. X 7-12ijl). Asci (and stalk cells) on stamens and calyx lobes
are intermediate in size. Similar size-differences in asci from leaves
and from fruits exist in Taphrina confu^a, T. farlowii, T. communis
and other species.

It is noteworthy that where collection dates are given the fungus
on leaves was collected later in the season than on fruits. This is in
accord with the writer's experience with Taphrina. confusa on P. vir-
giniana var. demissa in Colorado. Leaf curl of this host can be col-
lected long after the diseased fruits and inflorescences have been
shed.

Mix: Monograph of Genus Taphrina

139

Fig. 35. Asci (X900) of A, Taphrina padi; B, T. thomasii; C, T.

purpurascens.

140 The University Science Bulletin

No asci nor stalk cells as long as those reported by Ray (1939)
were seen in this study. A striking feature was the variability in
length of the stalk cell in different microscopic mounts, whether
from leaves, fruits, or floral parts.

80. Taphrina padi (Jacz.) Mix

Taphrina padi (Jacz.) Mix Trans. Kansas Acad. Sci. 50:77-83. 1947.
T. pruni Tulasne, Ann. Sci. Nat. 5 ser. Bot. 5:122-136. 1866, in part.
T. pruni Tul. var. padi Jaczewski, Pocket Key for the determination of
fungi. Part I. Exoascales. Leningrad. 1926.

Causing deformed fruits (pockets) on Primus padus L.

Mycelium intercellular.

Asci clavate, rounded at the apex, stalk cells variable, sometimes
widened at the base. Ascospores eight, round, ovate, or elliptic, fre-
quently budding in the ascus. (Fig. 35, A.)

Dimensions: Of asci, 26-46ijl X 8-13[x; of stalk cells, 8-26[jl X
7-lOtJL; of ascospores, 5-6[i X 4:-5\l.

Distribution: Europe.

Material examined: Sweden: Jiimtland, Storlien, Aug. 24, 1939,
A. J. M.; Lappland, Sulitolma near Lairo, July 29, 1938, T. Arwids-
son; Stockholm, Experimentalfaltet, July 5, 1939, C. Hammarlund
(type).

This fungus has long been considered to be identical with Taph-
rina pruni, though Sadebeck (1893) suggests that the fungi on
Pninus domestica and P. padus may not be the same. Jaczewski
(1926) separated the form on P. padus as a variety. Prunus padus
is so distantly related to P. domestica and P. insititia, that it seems
very unlikely that the two fungi are identical. Taphrina padi has a
distinctive morphology. It is clearly more closely related to Taph-
rina farlowii and to T. confusa than to T. pruni.

81. Taphrina confusa (Atk.) Giesenhagen

Taphrina confusa (Atk.) Giesenhagen, Flora 81:267-361. 1895.

Exoascus conjmus Atkinson, Bull. Torr. Bot. Club 21 :372-380. 1894.

T. unilateralis (Pk.) Mix, Univ. Kansas Sci. Bull. 24:10:151-176. 1936.

E. unilateralis Peck, New York State Mus. Nat. Hist. Ann. Rept. 51:267-

312. 1898.

T. cecidomophila (Atk.) Giesenhagen, I.e.

E. cecidomophilus Atkinson, I.e.

Deforming fruits and all floral parts of Prunus alabamensis Mohr,
P. virginiana L., P. virginiana L. var. demissa Torr., and P. virgin-
iana L. var. melanocarpa Sarg.

Mycelium intercellular.

Asci amphigenous on leaves, also covering surfaces of fruits and
floral parts, clavate, rounded at the apex; stalk cells variable in

Mix: Monograph of Genus Taphrina 141

size and shape, often irregular in outline. Ascospores eight, round,
ovate, or elliptic, frequently budding in the ascus. (Fig. 34, E-H).

Dimensions: Of asci, 23-53[x X 7-13[j,; of stalk cells, 6-27[J!. X
5-13[jl; of ascospores, 4-7[ji, X 3.5-6ijl.

Distribution: North America.

Material examined: P. alahamensis. Georgia: Pine Mountain,
May 12, 1936, J. H. Miller.

P. virginiana. Florida: Gainesville, April 17, 1936, G. F. Weber
(leaves). Massachusetts: Cape Cod, Sandwich, June 6, 1937, D.
H. Linder (leaves). Michigan: Glen Haven, Day State Park,
June 1, 1941, E. A. Bessey (leaves, fruits, and other floral parts).
Minnesota: Bear River, Big Bear Lake, June 25, 1947, A. J. M.
Nebraska: Dixon Co., Allen, May, 1940, R. W. Goss (leaves) ; Lin-
coln, June 3, 1940, L. B. Walker (fruits). New York: Brookton,
June, 1904, Fletcher (Cornell Univ. Dept. Plant Path. 20,000,
fruits); Etna, June, 1894, G. F. Atkinson (Atkinson Coll. 1192,
fruits deformed by insect larvae ; apparently type material of Exoas-
cus cecidomophilus) ; ibid., id. (two slides in Atkinson Collection,
numbered 1192) ; Evans Mills, June, C. H. Peck C'type No. 1" of
E. unilateralis, leaves) ; North Elba, June, 1897, id. ("type No. 2"
of E. unilateralis, leaves) ; Sempronius, June 5, 1937, W. W. Ray
(leaves and fruits) ; ibid., June 7, 1936, id. (fruits and other floral
parts). Nova Scotia: Pictou Co., Aug., 1910, L H. Crowell
(fruits). Ontario: London, June, 1893, J. Dearness (N. A. F. 2285,
fruits). Finland: near Mustiala, June, 1886, P. A. Karsten (F.
Eur. 3473, fruits).

P. virginiana var. demissa. California: Berkeley, May 15, 1937,
H. Earl Thomas (leaves) ; Mt. Shasta, Wagon Creek, June 7, 1941,
W. B. Cooke (Herb. W. B. C. 15506, leaves, young ovaries, and
other floral parts). Colorado: Boulder Canyon, El Vado, July 5,
15, 1943, A. J. M. (leaves) ; Cross Ruxton, July 5, 1906, F. E. and
E. S. Clements (Crypt. Form. Col. 311, leaves) ; Manitou, Mani-
tou Trail, July 13, 1895, L. H. Pammel (Herb. L. H. P. 114, leaves) ;
Ute Pass, slide No. 97 of Atkinson Collection (fruits). Idaho:
Farragut, May, 1944, R. H. Thompson (leaves and inflorescences).
Oregon: Eugene, May 30, 1930, J. R. Hansbrough (Herb. J. R. H.
612, leaves). Washington: Klickitat Co., Gulers, May 30, 1920,
J. S. Boyce (Herb. J. S. B. 494, leaves) ; Pullman, June 20, 1893, C.
V. Piper (F. Wash. 138, floral parts).

P. virginiana var. melanocarpa. Colorado: Fort Collins, June,
1935 (Bot. Dept. Colorado State Coll., leaves). Montana: Madi-

142 The University Science Bulletin

son Co., Madison River Canyon, June 3, 1914, H. M. Jennison
(Flor. Mont. 107, leaves).

The case for synonymy of Taphrina conjusa and T. imilateralis
has been well presented by Ray (1939). Numerous collections in
which leaves, fruits (or ovaries) and other floral parts are affected,
and in which the fungus is morphologically similar on all affected
parts, leave no doubt as to this synonymy. Peck (1898) gave meas-
urements for "Exoascus unilateralis" as: asci, 40-52[ji, X 13-16pL;
stalk cells, IS-lGpi broad and "about as long." Examination of
Peck's type specimens showed the asci to be 23-30[jl X 8-13i;,; stalk
cells, 8-18[;. X 7-lOpi. Obviously Peck's measurements are in error.

The fungus on Primus alahamensis with asci measuring 23-34tJL X
10-12[ji., stalk cells 8-10[j. X 9-llpi. is intermediate in size between
Taphrina farlowii and T. conjusa. It might well have been assigned
to T. farlowii or called a new species. Its host is apparently more
closely related to P. serotina than to P. virginiana. It is tentatively
placed with Taphrina conjusa because it deforms all floral parts, as
does that fungus.

The collection by Piper at Pullman, Washington, is interesting.
All floral parts, petals, sepals, stamens, and calyx-cup are enlarged
and bear asci, while the ovaries are prolonged into long conical
"pockets" 2-3 cm. long, with a curving beak, suggestive of Taphrina
andina {T. atkinsonii) .

Forms on Cherries: Laurocerasus

82. Taphrina thomasii Mix
Taphrina thomasii Mix, Trans. Kansas Acad. Sci. 50:77-83. 1947.

Causing witches' brooms on Prunus ilicijolia Walp.

Mycelium intercellular.

Asci hypophyllous, cylindric-clavate, rounded at the apex; stalk
cell somewhat variable, occasionally forked below, sometimes lack-
ing. Ascospores eight, round, ovate, or elliptic, frequently budding
in the ascus. (Fig. 35, B.)

Dimensions: Of asd, 20-35pi, X 6-lOpL (or if stalk cell lacking
32-38[j. X 6-lOtJi.) ; of stalk cells, 8-16[;, X 6-8[ji,; of ascospores, 4-7\i.
X 4-5;;..

Distribution: California.

Material examined: California: Santa Clara Co., Los Altos,
1943, H. Earl Thomas (type) ; ibid., May 8, 1947, id. (co-type).

In this species the stalk cell is cut off late and is sometimes lack-
ing in the mature ascus. The ascus emerges from the chlamydo-

Mix: Monograph of Genus Taphrina li'-i

spore, the thicker wall of the latter persisting around the stalk cell
or ascus-base.

HOST INDEX TO SPECIES OF TAPHRINA ON PRUNUS

Prunics alahamensis Mohr

Taphrina conjusa (Atk.) Gies.
Prunits americana Marsh.

Taphrina communis (Sadeb.) Gies.
Prunus angustijolia Marsh.

Taphrina communis (Sadeb.) Gies.
Prunus armeniaca L.

Taphrina armeniacae Georg. and Bad.
Prunus armeniaca L. var. ansu Maxim.

Taphrina mume Nish.
Prunus avium L.

Taphrina cerasi (Fkl.) Sadeb.
Prunus bcsseyi Bailey

Taphrina flavorubra Ray
Prunus cerasifera Ehrh. var. divaricata Bailey

Taphrina pruni Tul.
Prunus cerasus L.

Taphrina cerasi (Fkl.) Sadeb.
Prunus cerasus L. var. frutescens (Neilr.) Schneid.

Taphrina cerasi (Fkl.) Sadeb.
Prunus communis Arcang. var. amara Schneid.

Taphrina deformans (Berk.) Tul.
Prunus communis Arcang, var. didcis Schneid.

Taphrina deformans (Berk.) Tul.
Prunus domeslica L.

Taphrina pruni Tul.
Prunus cmarginata Walp.

Taphrina flectans Mix
Prunus fruiicosa Pall.

Taphrina cerasi (Fkl.) Sadeb.
Prunus hortidana Bailey

Taphrina comtnunis (Sadeb.) Gies.
Prunus hortulana Bailey var. mineri Bailey

Taphrina communis (Sadeb.) Gies.
Prunus humilis Bunge

Taphrina cerasi-microcarpae (Kuschke) Laubert
Prunus ilicifolia Walp.

Taphrina thomasii Mix
Prunus incisa Thunb.

Taphrina truncicola Kus.
Prunus insititia L.

Taphrina pruni Tul.
Prunus lanata Mack, and Bush.

Taphrina communis (Sadeb.) Gies.
Prunus mandshurica Koehne

Taphrina mume Nish.
Prunus maritima Marsh.

Taphrina communis (Sadeb.) Gies.
Prunus maximowiczii Rupr.

Taphrina truncicola Kus.
Prunus microcarpa C. A. Mey.

Taphrina cerasi-microcarpae (Kuschke) Laubert
Prunus microphylla (HBK) Gray

Taphrina mexicana H. and P. Syd.
Prunus mume Sieb. and Zucc.

Taphrina inume Nish.

144 The University Science Bulletin

Prunus munsoniana Bailey

Taphrina communis (S'adeb.) Gies.
Primus nigra Ait.

Taphrina communis (Sadeb.) Gies.
Prunus padus L.

Taphrina padi (Jacz.) Mix
Prunus pennsylvanica L.

Taphrina cerasi (Fkl.) Sadeb.
Prunus persica L.

Taphrina deformans (Berk.) Tul.
Prunus pseudocerasus Lindl.

Taphrina cerasi (Fkl.) Sadeb.
Prunus pumila L.

Taphrina flavorubra Ray
Prunus pumila L. var. susquehanae Jaeg.

Taphrina flavorubra Ray
Prunus salicina Lindl.

Taphrina communis (S'adeb.) Gies.
Pi-umis scrotina Ehrh.

Taphrina farlowii Sadeb.
Prunus serotina Ehrh. var. salicijolia Koehne

Taphrina andina Palm
Prunus serrulata Lindl. var. lannesiana Rehd.

Taphrina cerasi (Fkl.) Sadeb.
Prunus serrulata Lindl. var. spontanea Wils.

Taphrina cerasi (Fkl.) Sadeb.
Prunus s'ptnosa L.

Taphrina pruni Tul.
Prunus subcordata Benth.

Taphrina pruni-subcordatae (Zeller) Mix
Prunus subhirtella Miq.

Taphrina cerasi (Fkl.) Sadeb.
Prunus tomcntosa Thunb.

Taphrina cerasi-microcarpac (Kuschke) Laubert
Prunus umbellata Ell.

Taphrina communis (S'adeb.) Gies.
Prunus ussuriensis Koval. and Kost.

Taphrina pruni Tul.
Prunus Virginian a L.

Taphrina conjusa (Atk.) Gies.
Prunus virginiana L. var. demissa Torr.

Taphrina conjusa (Atk.) Gies.
Prunus virginiana L. var. melanocarpa Sarg.

Taphrina conjusa (Atk.) Gies.
Prunus yedoensis Matsum.

Taphrina cerasi (Fkl.) Sadeb.

Mix: Monograph of Genus Taphrina 145

VII. SPECIES ON ANACARDIACEAE

Rhus
83. Taphrina purpurascens Robinson

Taphrina purpurascens Robinson, Ann. Bot. 1:163-176. 1887.

Ascomyces deformans Berk. var. purpurascens Ellis and Everhart, North

American Fungi No. 1886.

Causing leaf -curl, with great enlargement and reddish-purple
coloration of leaves of Rhus copallina L., R. coriaria L., R. glabra
L., and R. typhina L.

Mycelium intercellular.

Asci hypophyllous or amphigenous, dumbbell-shaped, constricted
in the middle, rounded at the apex, rounded or truncate at the base,
which is sometimes wider than the apex. No stalk cell. Ascospores
eight, frequently budding in the ascus, round, ovate, or elliptic.
(Fig. 35, C.)

Dimensions: Of asci, 17-40[x X 9-17[ji. (width of constricted part
6-12[ji,) ; of ascospores, 3-6[x X 3-5[ji..

Distribution: North America, France, Sicily.

Material examined: Rhus coriaria. France: Mountpellier, May
13, 1905, G. Arnaud; ibid.. May 21, 1914, id.

Rhus copallina. Arkansas: Fayetteville, June 10, 1937; ibid.,
May 24, 1941, J. C. Dunegan and A.J. M.; Johnson Co., May 13,
1935, W. C. Amstein. Kansas: Neosho Co., 1 mi. E. of Morehead,
June 30, 1947, R. L. McGregor; Wilson Co., 3 mi. E. of Neodesha,
July 1, 1947, R. L. McGregor; Woodson Co., 3 mi. N. W. of Yates
Center, July 1, 1947, R. L. McGregor. Massachusetts: Manches-
ter, July, 1880, W. C. Sturgis (Econ. F. 120a) ; Pigeon Cove and
Magnolia, July 28, 1890, A. B. Seymour (Econ. F. 120b). New
Jersey: Newfield, June, 1886, Ellis and Everhart (N. A. F. 1886).
Virginia: 1935, S. A. Wingard.

R. glabra. Connecticut: New Haven, R. Thaxter (Farl. Herb.
3661).

R. typhina. Connecticut: New Haven, May, 1889, R. Thaxter;
ibid.. West Rock, id. (Farl. Herb. 1498) .

Material was distributed in Fungi Columbiana 4024, as Exoascu^
purpurascens (E. and E.) Sacc. on Rhu^ aromatica Ait., Lake
Huron, Ontario, Canada, Aug. 10, 1912, J. Dearness. Several speci-
mens of this collection have been seen and in none of them was a
fungus found. The reddish distortions of the leaves appear to be
the work of insects.

Montemartini (1940) reports this fungus as occurring on Rhu^
coriaria L. in Sicily.
10—3659

146 The University Science Bulletin

VIII. SPECIES ON ACERACEAE

Acer.
84. Taphrina acerina Eliasson

Taphrina acerina Eliasson, Bih. K. Svensk. Vetensk. Akad. Handl. 20:3:4:

3-6. 1895.

Exoascus acerinits (Eliass.) Saccardo, Sylloge Fungoi-um. 14:823.

Causing leaf-blight and (according to Eliasson) witches' brooms
of Acer platanoides L. Affected areas on leaves are yellow-brown,
watersoaked in appearance (papery when dry), extending from
margin in between veins, affecting part or all of the leaf.

Mycelium subcuticular.

Asci amphigenous, broad-cylindric, rounded or truncate at the
apex, stalk cell broader than the ascus. Ascospores eight, round,
ovate, or elliptic, often budding in the ascus. Ascus emerging from
the chlamydospore by rupture of the chlamydospore wall. (Fig.
36, A.)

Dimensions: Of asci, 12-23iJt. X 8-12pt,; of stalk cells, 7-13i/. X
10-20[j. ; of ascospores, 4-5[x X 3.5-5ijl.

Distribution: Norway, Sweden, Russia.

Material examined: Norway: Anst-Agder, M0glestne in Vestre
Moland, June 16, 1932, I. J0rstad. Sweden: Uppland, Uppsala,
Stafsund, June 28, 1893, A. G. Eliasson; Vastergotland, Mariestad,
June, 1911, N. Sylven.

85. Taphrina acericola Massalongo

Taphrina acericola Massalongo, Malpighia 8 :97-130. 1894.

Exoascus acericolus (Massal.) Saccardo, Sylloge Fungorum 11:430.

T. jaczewski Palm, Arkiv. Bot. 15:1-14. 1917.

E. confusus Jaczewski, Bull. Jard. Imp. Bot. St. Petersburg. 1:7-13. 1901.

Causing small (2-3 mm. diam.), well defined, definitely margined
spots, with a tendency toward extension and coalescence, on leaves
oi Acer campestre L.

Mycelium subcuticular.

Asci hypophyllous or amphigenous, short-cylindric to ellipsoid,
rounded at the apex; stalk cell short and broader than the ascus;
ascospores eight, round, ovate, or elliptic, often budding in the
ascus. Ascus emerging from the chlamydospore by rupture of the
chlamydospore wall. (Fig. 36, B, C.)

Dimensions: Of asci, 16-26[ji. X 6-13[x; of stalk cells, 4-13[/, X
8-17[j,; of spores, 4-5.5[Ji X 3.5-4.5[ji,.

Distribution: Italy, Caucasus.

Material examined: As T. acericola. Italy: Spiassi, May 28,

Mix: Monograph of Genus Taphrina

147

Fig. 36. A, asci, ascus emerging from chlamydospore of Taphrina
acerina; B. asci of T. acericola; C, asci, chlamydospores, and emerging
ascus of T. acericola {T. jaczewski) ; D. asci of T. sacchari; E, T. pseudo-
platani; F, T. bartholomaei. All X 900.

148 The University Science Bulletin

1895 (Ex. Herb. C. Massal). As T . jaczewski. Russia: Trancau-
casia, Kedanah, July, 1890 (part of type) ; Central Russia, June 13,
1913, N. Troussova (Herb. Inst. Mycol. ct Phytopath. Jaczewski).

Jaczewski (1901) distinguished Exoascus confusus from Taphrina
acericola by the fact that its mycelium is perennial in the branches
and induces witches' brooms. The type material shows only small
spots on leaves (quite like those caused by T. acericola) , but assum-
ing that Jaczewski is correct, the formation of witches' brooms
would be insufl&cient ground for separation of this fungus from T.
acericola, when the two are morphologically indistinguishable.

This fungus is very similar to Taphrina acerina, the asci of the
two forms being nearly alike in size and shape. In both fungi the
ascus emerges from the chlamydospore by rupture of the chlamydo-
spore wall. Both are reported as causing witches' brooms. It is
possible that they are the same species.

86. Taphrina sacchari Jenkins
Taphrina sacchari Jenkins, Jour. Wash. Acad. Sci. 28:8:353-358. 1938.

Causing small, round to irregular, brownish marginal spots, which
may coalesce to form blighted areas extending inward between main
veins, on leaves of Acer saccharum Marsh., and A. migrum Michx.

Mycelium subcuticular.

Asci hypophyllous, often scattered, in places close packed, short-
cylindric, rounded at the apex; stalk cells short and broad, some-
times broader than the ascus. Ascospores eight, round, ovate, or
elliptic, often budding in the ascus. (Fig. 36, D.)

Dimensions: Of asci, 13-30ijl X 6-13[jl; of stalk cells, S-lSpL X
8-20[jl; of ascospores, 3.5-6[i X 3-5ijl.

Distribution: eastern and central North America.

Material examined: Acer saccharum. Arkansas: Fayetteville,
May 2 and May 5, 1935, V. H. Young; ibid., June, 1935, J. C. Dune-
gan; ibid.. May 21, 1937, V. H. Young; ibid.. May 8, 1940, id.; ibid..
May 24, 1941, J. C. Dunegan and A. J. M.; Mountainburg, May 25,
1941, A. J. M. Kansas: Independence, May 11, 1944, G. W. Staf-
ford. Maine: Rockport, June 26, 1922, J. Achorn (U. S. D. A. Path,
and Myc. Coll. 68554). Pennsylvania: Williamsport, June 17,
1935, R. S. Kirby (from L. 0. Overholts).

Jenkins (1938) in describing Taphrina sacchari mentioned its
close similarity to T. acericola but considered that she had evidence
of a biological difference between the two. The morphological simi-
larity is indeed very close and it may be that the two species are
identical. They are here treated as separate species because of
differences in host relationships and geographic distribution.

Mix: Monograph of Genus Taphrina 149

887. Taiphrina bartholomaei nomen novum

Taphrina bartholomaei nomen novum.

T. aceris (Dearn. and Barth.) Mix, Univ. Kansas Sci. Bull. 24:10:151-176.

1936.

Exoascus aceris Dearn. and Barth. (Dearness, Mycologia 9:345-364. 1917.)

Causing small (ca. 5-10 mm. diam.), brown, unthickened, necrotic
spots on leaves of Acer grandidentatum Nutt.

Mycelium subcuticular.

Asci hypophyllous, epiphyllous, or amphigenous, short-cylindric,
rounded at the apex; stalk cell short, often broader than the ascus.
Ascospores eight, round, ovate, or elliptic, often budding in the
ascus. (Fig. 36, F.)

Dimensions: Of asci, 20-40[jl X S-lSfji.; of stalk cells, 7-17[j. X
7-17pt.; of ascospores, 4-5.5[jl X 3.5-4.5[ji..

Distribution: Utah.

Material examined: Utah: Salt Lake Co., Parley's Canyon,
June 29, 1915, Bartholomew and Garrett (F. Columb. 5018).

The name Exoascus aceris applied to this fungus by Dearness,
and the new combination made by Mix (1936a) are invalid, since
Fisch (1885) had described a fungus on Acer platanoides as Exoas-
cus aceris Linhart. This fungus was declared identical with Taph-
rina polyspora Sorok. by Johanson (1886).

In renaming this fungus it would have been desirable to honor
Dearness but that had been done by Jenkins (1939) in naming a
fungus on Acer ruhrum. The species is therefore named after Bar-
tholomew, who took part in its collection.

88. Taphrina pseudoplatani (Massal.) Jaap

Taphrina pseudoplatani (Massal.) Jaap, Ann. Mycol. 15:97-124. 1917.

Taphrina polyspora (Sorok.) Johans. var. pseudoplatani Massalongo, Bull.

Soc. Bot. Ital. 1892:197-199.

T. acericola (Massal.) var. pseudoplatani Massalongo, Malpighia 8:97-130.

1894.

Causing irregular, unthickened, gray to blackish spots (0.5 to
1 cm. diam.) with indefinite margin, on leaves of Acer pseudopla-
tanus L.

Mycelium subcuticular.

Asci hypophyllous, short-cylindric, rounded at apex, stalk cell
short, usually not wider than the ascus. Ascospores eight, round,
ovate, or elliptic, often budding in the ascus. (Fig. 36, E.)

Dimensions: Of asci, 13-26iji. X 8-13[x; of stalk cells, 6-lS[i X
8-13pL; of ascospores, 4-5. 5ix X 3.5-4.5[jl.

Distribution: Italy, Switzerland.

Material examined: Switzerland: near. Grindelwald, Aug. 3,

150 The University Science Bulletin

1910, 0. Jaap. No locality, date or collector (Herb. C. Massalongo
and Ex Herb. Bresadola in Bot. Mus. Stockholm) .

This fungus seems very close to TapJmna acericola, from which
it is distinguished by its narrower stalk cell. It is significant that
both fungi occur in Italy. Perhaps further study will show them to
be identical.

89. Taphrina polyspora (Sorok.) Johanson

Taphrina polyspora (Sorok.) Johanson, Ofvers. of K. Svensk. Vetensk.

Akad. Forhandl. 1885 :29-47. 1886.

Ascomyces polysq)orus Sorokin. Ann. Sci. Nat. 6 ser. Bot. 4:72. 1876.

Exoascus aceris Linhartj Fungi Hungarici (Fisch, Bot. Centbl. 22:126-127.

1885).

Causing small (ca. 1 cm. diam.) roundish spots or larger irregular
areas, blackish, unthickened, on leaves of Acer, orientale L. and A.
tataricum L.

Mycelium subcuticular.

Asci hypophyllous, short-cylindric, rounded at the apex, often
widened at the base forming a fairly definite foot, lacking a stalk
cell. Ascospores eight, round, ovate, or elliptic, budding at once and
filling the asci with small round to elliptic blastospores. (Fig. 37,
A.) _ _

Dimensions: Of asci, 26-60[jl X 8-23|j.; of ascospores, 4.5-5[ji. X
4-4. Spi; of blastospores, 2-4[x X 2-4[x, or smaller.

Distribution: Europe.

Material examined: A. orientale. Greece: Messenie, Selitsa,
Mt. Taygetos, April 24, 1908, R. Maire (Mission Bot. Orient, 1908,
3,200.). A. tataricum. Germany: Bavaria, Freising, Garden of
Lyceum, June, 1902, J. E. Weiss (Myc. Germ. 173). Russia: Pet-
ropolitana, Peterhof, A. Jaczewski (Cornell Univ. Dept. Plant Path.
17092). Sweden: Uppland, Uppsala, June, 1884, C. J. Johanson.

90. Taphrina letifera (Pk.) Saccardo

Taphrina letifera (Pk.) Saccardo, Sylloge Fungorum 10:67.

Ascomyces letifer Peck, New York State Mus. Nat. Hist. Ann. Rept. 40:

39-77. 1887.

Causing large, indefinitely margined spots on leaves or blighting
whole leaves of Acer spicatum Lam. Spots when fresh may be green
or reddened, but in dried specimens are black.

Mycelium intercellular.

Asci hypophyllous or amphigenous, broad-cylindric, rounded or
truncate at the apex; stalk cell short, frequently wider than the
ascus. Ascospores eight, round, ovate, or elliptic, often budding in
the ascus. (Fig. 37, B, C.)

Mix: Monograph of Genus Taphrina

151

Fig. 37. Asci (X 900) of A, Taphrina polyspora; B, C, T. letifera; D, E,

T. darken.

152 The University Science Bulletin

Dimensions: Of asci, 14-35[Ji. X 8-18[;,; of stalk cells, 6-13[jl X
8-25[j,; of spores, 4-5[x X 4-5!;,.

Distribution: eastern North America.

Material examined: Massachusetts: Mt. Wachusett, June 12,
1936, G. D. Darker. Minnesota: Bear River, Big Bear Lake, June
28, 1947, A. J. M. New York: Elizabethtown, June, 1886, C. H.
Peck (part of type) ; Labrador Lake, near Apulia, June 25, 1940,
A. J. M. North Carolina: Nantahala Gorge, May 29, 1941, id.

Peck's measurements of this fungus are in decimal fractions of
inches. When converted into millimeters they become: Asci, 40-50[j(.
X 15-20pi,. Examination of Peck's type material shows these meas-
urements to be wrong. They would not be so far wrong if Peck
had been considering the whole ascus, including the stalk cell. He
makes no mention of the stalk cell.

Jenkins (1938) gives chmensions of asci as 36-50[jl X 14-20tJt., of
stalk cells as 14-20pL X 17-25pt. Careful study of the type material
as well as of other specimens mentioned above has revealed no asci
so large as these. Asci on the lower surface of the leaf are regularly
smaller than those on the upper.

Peck spelled his species name ''letifer." Saccardo inserted an "h"
in renaming the fungus Taphrina lethifera. As "letifer" is good
Latin, Saccardo's change was unnecessary.

Jenkins and Ray (1940) have assigned the fungus collected by
them on Acer spicatum at Labrador Lake to Taphrina dearnessii on
the basis of similarity of host lesions and size of asci. Material col-
lected at Labrador Lake by the writer agrees in all respects with
Taphrina letifera. It may be that Taphrina dearnessii and T. leti-
fera will prove to be one and the same species ; the two fungi were
found abundantly in the same locality (Bear River, Minn.) in 1947.
In any event convincing evidence that two distinct species of Taph-
rina occur on Acer spicatum is lacking.

91. Taphrina darken Mix
Taphrina darken Mix, Trans. Kansas Acad. Sci. 50:77-83. 1947.

Causing small brown necrotic spots on leaves of Acer circinatum
Pursh.

Mycelium intercellular.

Asci amphigenous, broad-cylindric, rounded or truncate at the
apex; stalk cell short, often wider than the ascus; ascospores eight,
round, ovate, or elliptic, frequently budding in the ascus. (Fig. 37,
D, E.)

Mix: Monograph of Genus Taphrina 153

Dimensions: Of asci, 16-3S[k X 8-13[j.; of stalk cells, 5-12[x X
10-17[jl; of spores, 4-4.5[;, X 3.5-4[;,.

Distribution: Oregon.

Material examined: Oregon: Oregon Caves, head of Limestone
Creek, Aug. 15, 1929, G. D. Darker (Herb. Arnokl Arboretum 5077).

92. Taphrina nikkoensis Kusano

, Taphrina nikkoensis Kusano, Bot. Mag. Tokyo. 21 :65-67. 1907.
Exoascus nikkoensis (Kus.) Saccardo, Sylloge Fungorum 22:763.

Causing small (ca. 5 mm. diam.) round to irregular, yellowish,
unthickened spots, or larger blighted areas on leaves of Acer dia-
bolicum K. Koch. var. purpurascens Rehd.

Mycelium subcuticular.

Asci hypophyllous, cylindric, with rounded apex, rather variable in
«ize and shape; stalk cell no wider than the ascus- Ascospores eight,
round, ovate, or elliptic, often budding in the ascus. (Fig. 38, A.)

Dimensions: Of asci, 20-50iJi, X 8-13[;.; of stalk cells, 8-15[j, X
7-13[ji,; of spores, 4-6ii, X 3.5-5tJL.

Distribution: Japan.

Material examined: Japan: Nikko, May 27, 1906, S. Kusano
(Received from Kusano, presumably part of type) .

93. Taphrina dearnessii Jenkins

Taphrina dearnessii Jenkins, Jour. Washington Acad. Sci. 29:222-230. 1939.

Causing small, well defined dark brown to black, unthickened
spots, sometimes coalescing to involve large areas of the leaf-blade,
on leaves of Acer rubrum L.

Mycelium subcuticular.

Asci amphigenous, short-cylindric, rounded or truncate at the apex,
stalk cell short, often broader than the ascus. Ascospores eight,
round to elliptic, often budding in the ascus. (Fig. 38, B.)

Dimensions: Of asci, 17-35[jl X 8-17[jl; of stalk cells, 6-13[x X
10-23[ji.; of ascospores, 4.5-6ij. X 3-5^;..

Distribution: eastern North America.

Material examined: Michigan: Indian River, July 1, 1947, A. J.
M.; between Isabella and Cooke, July 1, 1947, id. Minnesota:
Bear River, Big Bear Lake, June 29, 1947, A. J. M.; Janet Lake, 28
mi. N. of Floodwood, July 1, 1947, id. New York: Ithaca, Ring-
wood Preserve, June 11, 1940, A. J. M. North Carolina:
Nantahala Gorge, May 29, 1941, id. Virginia: Hahfax Co.,
Nathahe, May 13, 1935, E. R. Mickle. Quebec: Davidson, June,
1938. 1. L. Connors.

154

The University Science Bulletin

Fig. 38. Asci (x 900) of A, Taphrina nikkoensis; B, T. dearnessii; C,
T. carveri; D, T. sebastianae ; E, T. aesculi.

Mix: Monograph of Genus Taphrina 155

94. Taphrina carveri Jenkins
Taphrina carveri Jenkins, Jour. Washington Acad. Sci. 29:222-230. 1939.

Causing small, well defined black spots on leaves of Acer sac-
charinuvi L.

Mijcelium subcuticular.

Asci hypophyllous, short-cylindric, rounded or truncate at the
apex; stalk cells short, often broader than the ascus. Ascospores
eight, round, ovate or elliptic, often budding in the ascus. (Fig.
38, C.)

Dimensions: Of asci, 23-53iji, X 8-17tJ.; of stalk cells, 4-15pL X
10-221JI,; of spores, 4.5-6[x X 3-5iji,.

Distribution: Alabama, Missouri.

Material examined: Alabama: Tuskegee, April 30, 1897 (Ellis
Herb. 17918). Missouri: Lutesville, May 30, 1941, A. J. M.; i6id.,
June 3, 1942, L. Engelhart.

This fungus is very close indeed to Taphrina dearnessii on Acer
rubrum. Perhaps the two species are identical.

HOST INDEX TO SPECIES OF TAPHRINA ON ACER

Acer canipestre L.

Taphrina acericola Massal.
Acer circinatum Pursh.

Taphrina darkeri Mix
Acer diabolicum K. Koch. var. purpurascens Rehd.

Taphrina nikkoensis Kus.
Acer grandidentatum Nutt.

Taphrina bartholomaei (Deam. and Earth.) Mix
Acer nigrum Michx.

Taphrina sacchari Jenkins
Acer orientate L.

Taphrina polyspora (Sorok.) Johans.
Acer platanoides L.

Taphrina acerina Eliass.
Acer pseudoplatanus L.

Taphrina pseudoplatani (Massal.) Jaap
Acer rubrum L.

Taphrina dearnessii Jenkins
Acer saccharum Marsh.

Taphrina sacchari Jenkins
Acer saccharinum L.

Taphrina carveri Jenkins
Acer spicatum Lam.

Taphrina letijera (Pk.) Sacc.
Acer tataricum L.

Taphrina polyspora (Sorok.) Johans. '

156 The University Science Bulletin

IX. SPECIES ON HIPPOCASTANACEAE

Aesculus

95. Taphrina aesculi (Patt.) Giesenhagen

Taphrina aesculi (Patt.) Giesenhagen, Flora 81:267-361. 1S95.

Exoascus aesculi Patterson, Bull. Lab. Nat. Hist. Univ. Iowa 3:89-135.

1895.

Causing small to large, round to irregular, yellow-brown blighted
areas on leaves of Aesculus calif ornica Nutt. Affecting whole shoots
and shoot-systems. Specimens seen have not been true witches'
brooms.

Mycelium intercellular, in leaves and twigs.

Asci hypophyllous, epiphyllous, or amphigenous, rounded at the
apex, stalk cell short, sometimes wider than the ascus. Ascospores
eight, round, ovate, or elliptic, often budding in the ascus. (Fig.
38, E.)

Dimensions: Of asci, 20-38[ji, X 8-15[J^; of stalk cells, 7-13[ji. X
10-23i;,; of spores, 4-7ijl X 4-5.5!x.

Distribution: California.

Material examined: California: Glenn Co., near Elk Creek, May
15, 1938, H. Earl Thomas; Hopland, May 13, 1936, id.; Marin Co.,
Muir AVoods, May 25, 1929, L. Bonar; Palo Alto, San Francisquito
Creek, April 25, 1940, R. H. Thompson; ibid., May 1, 1940, id.;
ibid., May 13, 1941, id.

Mix: Monograph of Genus Taphrina 157

X. SPECIES ON COMPOSITAE

Sebastiana

96. Taphrina sebastianae (Sadeb.) Jaczewski

Taphrina sebastianae (Sadeb.) Jaczewski, Pocket Key for Determination

of Fungi. Part I. Exoascales. Leningrad. 1926.

Exoascus sebastianae S'adebeck, Ber. Deutsch. Bot. Ges. 22:119-133. 1904.

Causing lesions whose nature cannot be determined from extant
material (because of the presence of a rust-fungus) on leaves of
Sebastiana ypanemensis (Mill.) Arg.

Mycelium of unknown habit.

Asci cylindric, rounded at the apex, stalk cells flattened or wedge-
shaped below. Ascospores eight, round, ovate, or elliptic, often
budding in the ascus. (Fig. 38, D.)

Dimensions: Of asci, 26-40ijl X 13-18[ji.; of stalk cells, T-lSpi, X
13-18pi.; of blastospores, 4-5.5[/, X 2-4[x.

Distribution: Brazil.

Material examined: Brazil: Santa Catharina, Tubarao, Nov.
1889, E. Ule [three collections: 1. E. Ule, Fungi botanicum Beroli-
nense 1496, Type. 2. Ex. Herb. Sydow (in Bot. Mus Stockholm).
3. Herbarium Brasiliense 1495 (in Farl. Herb.) ].

Sadebeck (1904), in describing this fungus, recorded a remarkable
polymorphism of asci. All of the curious asci described and figured
by him were apparently teliospores of a rust. They occur in all
states: immature, mature, and germinating. The rust is abundant
in all specimens examined and in the type specimens no asci are to
be found. The asci were seen best in the specimens in the Stock-
holm Museum. They are typical Taphrina-asci as described above.
Further collections of this fungus would be very desirable.

The labels of the type-collection and of the specimens in the
Farlow Herbarium read: "Exoascus Sebastianae Sadeb. n. sp.,
Uredo Sebastianae Wint. n. sp."

158

The University Science Bulletin

XI. SPECIES ON ZINGIBERACEAE

Curcuma, Globba, Hedychium, Zingiber

97. Taphrina maculans Butler

Taphmia maculans Butler, Ann. Mycol. 9:36-39. 1911.

Causing small (up to 5 mm. diam.) yellow-brown, unthickened

spots, (very numerous and crowded) on leaves of Curcuma amada

Roxb., C. angustifolia Roxb., C. longa L., Hedychium acuminatum

Wall, Zingiber casumunar Roxb., Z. mioga Rose, Z. berumbet Rose.

Fig. 39. A, ascogenous cells; B, asci of Taphrtna maculans; C, asci;
D, ascogenous cells of T. linearis. E-H, mycelium of T. lineaiis growing
in the epidermal wall. All X 900.

Mix: Monograph of Genus Taphrina 159

Mycelium growing within epidermal walls and within radial walls
between cells of epidermal and subepidermal layers, sometimes form-
ing fiat plates of hyphae, occasionally sending haustoria into host
cells. (Fig. 39, A, C.)

Asci epiphyllous, clavate, rounded or truncate at the apex; stalk
cell single or divided by septa into two or three cells; ascospores
eight, round, ovate, or elliptic, sometimes budding in the ascus.

Dimensioxis : Of asci, 20-36[jl X 6-lOpL; of stalk cells, 10-17[jl X
5-l[i; of ascospores, 4-6. 5[x X 2,-3.5\}..

Distribution: India, Japan.

Material examined: Curcuma amada. India: Sahranpur, Gov-
ernment Bot. Gardens, Nov. 17, 1903, W. Gollan.

C. angustifolia. Himalaya: Kumaon, Bageshwar, June 13, 1907,
Inayat Khan.

C. longa. Bihar: Pusa, Aug., 1916, M. Taslim. Bombay Pres.:
Hatkalangda, Dec. 15, 1910, G. S. Kulkarni.

Hedychium acuminatum. India: Nainital and Musscorie, Oct.
6, 1929, J. H. Mitter, A. K. Mitra, V. B. Singh.

Zingiber casumunar. India: Bengal, Rangpur, March 13, 1909,
S. N. Mitra.

Z. mioga. Japan: Mutsu, Korekawa, Oct. 1, 1913, M. Miuma
(Herb. Sydow, Bot. Mus. Stockholm).

Z. zerumbet. India: Bihar, Muzaffarpur, Awapur, Oct. 18, 1910,
Jamal bux.

Most of these specimens are immature, and only rarely may ma-
ture asci be found.

98. Taphrina linearis Sydow and Sydow
Taphrina linearis Sydow and Sydow, Ann. Mycol. 12:545-576. 1914.

Causing small, elongate, red-brown spots or streaks on leaves of
Globba marantina L.

Mycelium growing within the outer epidermal wall and within
radial walls between cells of epidermal and hypodermal layers.

Asci clavate, rounded or truncate at the apex, each provided with
one or a few stalk cells. (Fig. 39, B, D-H.)

Dimensions: Of asci, given by the authors as 25[jl X 7-9[x; a few
young asci seen by the writer measured 13-20[jl X 6-8iJi,; of asco-
spores, 4-5[jL X 3.5-4[ji,.

Distribution: Philippine Islands.

Material examined: Philippine Islands: Luzon, Bulacan, near
Angat,.Sept. 15, 1913, M. Ramos (F. Exot. Exs. 422. Three copies
seen).

This material is, for the most part, immature.

160 The University Science Bulletin

XII. NONVALID AND EXCLUDED SPECIES
A. Described Species

1. Exoascus anomalus Saccardo, Sylloge Fungorum 8:820.
This is Ascocorticiimi anomalum Schroet.

2. Taphrina candicans Saccardo, Michelia 1:117-132. 1880.

This "fungus" was described by Saccardo as a doubtful species of
Taphrina occurring on Teucrium chamaedrys L. A specimen exists
in the Botanical Museum at Padua, and duplicates were issued (col-
lected at Lyon, 1884) under this name as Roumeguere, Fungi select!
exsiccati 4499. The species was excluded from the genus by Gies-
senhagen (1895) and the specimen is said by Jaczewski (1926) to
show the effect of mite-injury and no fungus.

3. Taphrina cissi Zollinger, Natur. et Genneeskund. Arch. Nederlands Indie
1:372-405, 2:1-19, 200-273, 563-587, 3:51-92. 1847.

This fungus was described as occurring on Cissus varius. A rec-
ord and description of it is to be found in the Botanical Museum at
Buitenzorg, but no specimen is known. It is not mentioned in the
literature since Zollinger except for a copy of Zollinger's description
in Saccardo, Sylloge Fungorum 10:68.

4. Ascomyces fulgens Cooke and Harkness, Grevillea 9:6-9. 1880.
Exoascus fulgens (Cke. and Harkn.) Saccardo, Sylloge Fungorum 8:820.

This "fungus" was described by Cooke and Harkness as occurring
on leaves of Arctostaphylus pungens HBK. It was later pronounced
by Harkness (1886) to be an "aphidian gall."

5. Exoascus marginatus Lambotte and Fautrey. (Roumeguere, Fungi Exsicati
Praecipue Gallici, 6228.)

This was described as occurring on leaves of Crataegus oxyacan-
tha L. The specimens, collected Aug. 1892 by F. Fautrey, show
only injury due to mites (Erineum).

6. Taphrina rhaetica Volkart, Ber. Deutsch. Bot. Ges. 21:477-481. 1903.
This fungus, described by Volkart as occurring on leaves of Crepis

blattarioides Vill., was transferred by Juel (1902) to the genus
Taphridium of the Protomycetaceae becoming Taphridium volkartii
(Volkart) Juel.

7. Taphrina umbelliferarum Rostrup, Bot. Tidskr. 14:230-243. 1885.

The fungus, described by Rostrup as occurring on Heracleum
sphondylium L. and Peucedanum palustre Moench., was trans-
ferred by Juel (1902) to the genus Taphridium, becoming Taphri-
dium umhellij erarum (Rostr.) Lagerh. and Juel.

Mix: Monograph of Genus Taphrina 161

8. Taphrina oreoselini Massalongo, Nuov. Giorn. Bot. Ital. 21:422-423. 1889.
Occurring on Peucedanum oreoselinum Moench., this fungus is

apparently identical with the preceding and was renamed by Juel
(1. c.) Taphridium umhellijerarum.

9. Taphnua githaginis Rostrup, Vidensk. Meddel. Natursk. Foren. Kj0ben-
havn. 1890:246-264. 1890.

This fungus, on Agrostemma githago L., from its description
clearly belongs to Taphridium. Neger (1905) proposes its transfer
to that genus, without, however, actually renaming it. Specimens
have not been seen.

10. Exoascus avellanae Connold, Plant Galls of Great Britain. London. 1909.
ExoasciLS coryli Lemee, Bull. Soc. Hort. del Orne, Alencon. 1917.

No fungus having been seen in association with this witches'
broom of Corylus avellana, Saccardo (Sylloge Fungorum 24:1301)
declared this binomial to be a nom^n nudum.

11. Taphrina randiae Rehm, Hedwigia 40:170. 1901.

This species, occurring on Randia sp., was named and described
from a specimen collected at Serra Orgaos, Brazil, by E. Ule. It
was reported as causing elliptic (1-2 cm. X 1.5-1 cm.) black spots
thickened above. The asci were said to measure 50[jl X 25^^, to lack
a stalk cell, and to be polysporous. A portion of the type-specimen
was received from the Berlin Botanical Museum as a gift. It con-
sisted of part of a leaf which had been cut in two, the cut passing
through a thickened black spot which appeared as described above.
This spot contained acervuli but no asci. No mycelium character-
istic of Taphrina could be found. The name Taphrina randiae thus
becomes a nomen dubium. No attempt was made to identify the
imperfect fungus.

12. Exoascixs uleanus F. Hemmings, Hedwigia 43:79-95. 1904.

This species has been fully discussed earlier in the section dealing
with forms on ferns. As pointed out there no identifiable fungus can
be found, either in the type specimen or in duplicates. The binomial
becomes a nomen dubium.

13. Exoascus theobromae Ritzema Bos, Tijdschr. Plantenziekt. 6:65-90. 1900.
In this paper Ritzema Bos described a witches' broom of Theo-

broma cacao L. occurring in Surinam. He stated that he found a
few asci present, but could not, on account of the state of the
material describe the fungus. Nevertheless he named it as above.
Went (IQO'f) studied the same materiaf, finding mycelium present

11—3659

162 The University Science Bulletin

but no asci, and was unable to decide whether the witches' broom
was due to a species of Taphrina. Subsequently Faber (1908) de-
clared Exoascus theobromae to be a nomen nudum.

According to Cook (1913) the witches' broom of cacao, said by
Ritzema Bos to be caused by Exoascus theobromae, was later as-
cribed by Van Hall and Dorst to C olletotrichum luxificum.

14. Taphrina bussei Faber, Arb. K. Biol. Anst. Land imd Forstw. 6:385-395.
1908.

This fungus was described by Faber as causing a witches' broom on
Theobroma cacao L. The disease was found in Kamerun, and was
held to be distinct from the witches' broom of Surinam (see above).
The fungus was said to have asci 15-17[x X Sp^ and spores 2.5pi. X
1.7[L. No mention was made of a stalk cell but the asci shown in
Faber's figure seem to be borne on the ends of septate hyphae. The
fine mycelium described and figured is intracellular.

This fungus has not received further mention in the literature.
Dr. von Schoenau, then chief curator of the Munich Botanical
Museum, wrote in 1938 that he had been informed by Faber that
the type-material of Taphrina bussei had been placed in the her-
barium of the Biologische Reichsanstalt fiir Land-und Forstwist-
schaft in Berlin, but that a thorough search instituted there in re-
sponse to his inquiry had failed to reveal it.

According to Ludwigs (1934), this witches' broom (which should
be rather called "overbranching") is due to adverse environmental
factors and not to any parasite.

B. Species named but never described

1. Exoascus quercus-lobatae Mayr, Die Waldungen von Nordamerika etc.
Munich. 1889.

This name, unaccompanied by any description, was given to a
supposed fungus causing a witches' broom on Quercus lobata Nee.
It is clear from Mayr's account that he never saw any fungus and
the cause of the witches' broom remains unknown. Taphrina
caerulescens does occur on this host.

2. Taphrina jagi Lamb. (See account given earlier in this paper.)

3. "Exoascus or Taphrina" on Fuschia. Silo Leopoldo, Rio Grande do Sul.
1905. Rick, Fungi Austro Americani 78.

Three packets of this collection are in the Farlow Herbarium, two
of them from the herbarium of F. Theissen. No Taphrina is present.

Mix: Monograph of Genus Taphrina 163

4. "Exoascus Taphrina in Myrtacea." Sao Leopoldo, Rio Grande do Sul,
1929. Rick, Expeditions in Brazil. In Farlow Herbarium.

These specimens have an appearance suggesting mite injury. No

Taphrina is present.

5. "Exoascus Tradescantiae ," Sao Leopoldo, Rio Grande do Sul. Rick, Ex-
peditions in Brazil. In Farlow Herbarium.

Acervuli are present, but no Taphrina.

6. "Exoasciis Symploci E. and E." on Symplocus tinctoria, Coopolis, Missis-
sippi, April 24, 1898. S. M. Tracy. Fadow Herbarium, 5326, Herb. S. M.
Tracy.

Specimens are of greatly deformed structures, perhaps fruits. A
fungus is present but is not determinable.

7. "Taphrina selaginellae P. Hennings ad int." on Sclaginella menziesii Hook,
and Gr., Hawaii: Molokai. Pali of Kalanpapa from Kalae. W. Hillebrand.
In Mus. Bot. Beriin (in 1938).

This fungus is mentioned by Laubert (1928). Specimens ob-
tained from the Berlin Museum consisted of one healthy and one
diseased shoot. The latter was dwarfed, with closely appressed,
partially developed scales. It was pale green to brownish and bore
a number of small brown spots covered with a white bloom. The
fungus present was a Hyphomycete with spores about the size of
those of Taphrina. No asci could be found and the slender hyphae
were intracellular.

8. "Exoascus sp., Taches bullaties sulres feuilles de Alnus acuminata o' A.
ferruginea are Carrio Amon a' San Jose." Jan., 1913. A. Tonduz. Mus.
Nac. de Costa Rica 164. In Farlow Herbarium.

Perhaps insect injury. No Taphrina.

164 The University Science Bulletin

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bei Exoasceen. Forsch. auf d. Geb. d. Pflanzenkr. und d. Immun. im
Pflanzenr. 3:139-176.

THE UNIVEESITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. I] April 20, 1949 [No. 2

A Preliminary Account of the Herpetology of the State of

San Luis Potosi, Mexico

Edward H. Taylor

Abstract: A herpetological collection belonging to the Louisiana State
University, acquired through the efforts of Mr. and Mrs. Robert Newman and
Charles R. Shaw in the State of San Luis Potosi, Mexico, is reviewed in some
detail. Two amphibians, ten lizards, and twenty-three snakes are added to
the known fauna. One new snake genus, Schmidtophis, and the following new
species are described: Xenosaurus nevmianorum, a lizard of the family Xeno-
sauridae; and three colubrid snakes, Schmidtophis rubriventris, Rhadiriaea
marcellae and Tantilla sham.

Under the immediate direction of Mr. George Lowery, the Uni-
versity of Louisiana has for several years been pursuing a study of
the avian fauna of the state of San Luis Potosi, Mexico. Several
field parties have been stationed there at various times, and while
their primary interests have been directed toward the collecting of
birds, nevertheless numerous specimens of other vertebrate groups
have been taken.

The most recent field party (1946-1947), consisting of Mr. Robert
J. Newman, his wife, Marcella Newman, and Mr. Charles R. Shaw,
were requested to obtain a representative lot of the herpetological
fauna. This task fell largely to Mrs. Newman, who succeeded in
collecting or acquiring more than 300 specimens. Aside from these,
there are thirty-five specimens taken by Mr. Robert J. Newman and
forty-four by Mr. Charles R. Shaw.

The collection is of especial importance, since the San Luis Potosi
area had been largely neglected by earlier collectors, and in the re-
cent herpetological explorations of Dr. Hobart M. Smith and me in
Mexico no intensive collecting was done in any part of this state.
Our journeys, however, have taken us across the eastern part of the
state several times ; once across the western part, and on one trip I

(169)

170

The University Science Bulletin

journeyed from the eastern part through the central plains area to
the capital. Thus, a number of specimen records were obtained.

Three Mexican biotic provinces (delineated by Dr. Hobart M.
Smith on the basis of herpetological data) include portions of the
state. The Austro-central, which takes in the drier, western two-
thirds of the state; the Austro-oriental, which includes most of the
highlands of the eastern part; and a small lowland eastern area be-
longing in the Veracruzian province.

The known herpetological fauna of the state is listed herewith:

Snakes
Leptotyphlops humilis tenuiculus
Leptotyphlops myopicus

myopicus
Constrictor constrictor imperator
Conopsis nasus

Diadophis regalis laetus
Drymarchon corais erebennus
Drymobius margaritijerv^

margaritijerus
Elaphe chlorosoma
Geophis latijrontalis
Heterodon nasicus kennerlyi
Hypsiglena ochrorhyncha

ochrorhyncha
Hypsiglena ochrorhyncha janii
Lampropeltis mexicana
Leptodeira annulaia

septentrionalis
Leptodeira maculata
Thalerophis mexicanus

mexicanus
Masticophis flagellum lineatulus
Masticophis flagellum testaceus
Masticophis mentovarius

mentovarius
Masticophis taeniatus australis
Masticophis taeniatus ruthveni
Oxybelis potosiensis
Pituophis deppei deppei
Pseu^tes poecilonotus argus

Lizards
Crotaphytus collaris collaris?
Ctenosaurus acanthura
Sceloporus cautus
Sceloporus ferrariperezi

melanogaster
Sceloporus goldmani

Rhadinaea crassa
Rhadinaea gaigeae
Salvadora lineata
Spilotes pullatus mexicanus
Tantilla atriceps
Tantilla deviatrix
Tantilla rubra
Tantilla wilcoxi rubricata
Toluca lineata lineata
Tropidodipsas sartorii sartorii
Matrix rhombifera btanchardi
Storeria dekayi texana
Storeria dekayi temporalineata
Storeria hidalgoensis
Storeria storerioides
Thamnophis eques cyrtopsis
Thamnophis macrostemma

megalops
Thamnophis melanogaster

canescens
Thamnophis sauritus proximus
Thamnophis scalaris scaliger
Micrurus fitzingeri

m,icrogalbineus
Bothrops atrox asper
Crotalus atrox
Crotalus lepidus lepidus
Crotalus molossus nigrescens
Crotalus scutulatus scututatv^
Crotalus triseriatus triseriatus

Sceloporus jarrovii minor
Sceloporus microlepidotus

disparilis
Sceloporus parvus parvus
Sceloporus parvus scutulatus
Sceloporus spinosus spinosus

Taylor: Herpetology of San Luis Potosi, Mexico 171

Sceloporus variabilis variabilis
Phrynosonia modestum
Phrynosoma cornutum
Phrynosoma orbiculare var.
Holbrookia texana
Ameiva undulata podarga

Turtles
Kinosternon integrum
Terrapene mexicana

Amphibians

Diemictylus kallerti
Bolitoglossa rufescens
Bolitoglossa platydactyla
Chiropterotriton multidentata
Scaphiopus couchii
Scaphiopus multiplicatus
Bujo cognatus
Bujo horribilis
Bujo valliceps
Bujo punctatu^

Tomodactylus macrotympanum
Syrrhophuji giittilatus
Syrrhophus latodactylus
Syrrhophus cystignathoides

Cnemidophorus gularis gularis
Cncmidophorus perplexus
Barisia ivibricatus ciliaris
Anelytropsis papillosus
Eumeces lynxe lynxe
Eumeces tetragrammus

Gopherus berlandieri

Syrrhophus campi
Leptodactylus melanonotus
Leptodactylus labialis
Eleutherodactylus latrans
Eleutherodactylu^ rhodopis
Acrodytes spilomma
Smilisca baudinii baudinii
Hyla eximia
Hyla picta
Hyla staujferi
Hyla arenicolor
Hyla miotympanum
Hypopachus cuneus cuneus
Rana pipiens

The present collection contains numerous species that here are
traced to this area for the first time. The following represent new
records for the State of San Luis Potosi.

Pseudoeurycea bellii
Eleutherodactylus hidalgoensis
Anolis sericeus
Aiiolis petersii

Corythophanes hernandesii
Laemanctus serratu^
Crotaphytus collaris baileyi
Holbrookia maculata approximans
Sceloporus jarrovii immucronatus
Sceloporus serrijer plioporus
Sceloporus olivaceus
Gerrhonotus liocephalus loweryi
Adelphicos quadrivirgatus

Quadrivirgatus
Ficimia streckeri
Geophis multitorques
Ninia diademata plorator

Rhadinaea decorata
Dryadophis melanolomus

veraecrucis
Drymarchon corais erebennus
Pituophis deppei jani
Lampropeltis trianguluvi arcijera
Lampropeltis iriangulum

polyzona
Elaphe flaviruja flaviruja
Elaphe laeta laeta
Pliocercv^ laticollaris
Thamnophis phenax phenax
Thamnophis rnarciana
Coniophanes imperialis imperialis
Coniophanes fissidens proterops
Trimorphodon tau
Bothrops nummijer

172 The University Science Bulletin

There is a possibility that the species formerly reported from the
State as Crotaphytus collaris collaris is identical with the one here
referred to C. collaris baileyi.

The forms described as new are:

Xenosaurusnewmanorum Rhadinaea marcellae

Schmid'tophis Tantillashawi

Schmidtophis rubriventris

A few specimens in the collection will be treated at a later time.

The larger part of the specimens were collected in the area about
the town of Xilitla which lies in the southeastern part of the State of
San Luis Potosi a few miles from the border of the State of Hidalgo
and equally close to the border of Queretaro. Some of the moun-
tains in this region exceed an elevation of 7,000 feet. A few of the
specimens come from this elevation on Cerro Coneja. The greater
part of the specimens, however, are from elevations between 2,000
and 3,000 feet.

Mrs. Newman was successful in interesting the townspeople in the
collection and many specimens were brought to her by farmers,
school children and others. This procedure made it almost impossi-
ble to obtain accurate elevation data. The surprising character of
the collection is the far greater proportion of snakes to the am-
phibians and lizards than is usual in such a collection.

CAUDATA

Only one species of salamander, Bolitoglossa platydactyla
(Gray), is represented in the collection by a numbered specimen.
From the stomachs of certain snakes in the collection four other
species have been taken, but it has not been possible, in all cases, to
ascertain the complete identity of these.

Three of these came from snakes of the genus Thamnaphis.
These are Thamnophis scalaris scaliger, T. marciana, and T.
phenops phenops. The fourth specimen came from the stomach of
an aberrant rattlesnake referred to Crotalus triseriatus triseriatus.

Four species of salamanders have previously been found within
the state limits. These are Diemyctylus kallerti Wolterstorff, Bol-
itoglossa rufescens (Cope), Bolitoglossa platydactyla (Gray), and
Chiropterotriton multidentata (Taylor).

Diemyctylus kallerti Wolterstorff

Diemyctylus kallerti Wolterstorff, Abh. Ber. Mus. Magdeburg, vol. 6, 1930, pp. 147-149,
pi. 3, fig. 1; text figs. 12, 13. (Type locality, Tampico, Veracruz.)

This species, known from Tampico, Veracruz, and from Villa
Juarez, San Luis Potosi, has remained very rare in collections. A

Taylor: Herpetology of San Luis Potosi, Mexico 173

specimen, consisting of a head, more or less intact, together with the
partly digested remains of the body and limbs, was recovered from
the stomach of a snake, Thamnophis marciana (Baird and Girard),
which was taken 6.5 miles east of Sabanito, San Luis Potosi.

The generic identification was made on the characters of the head,
especially the teeth, choanae, and on the character of the vertebrae.
The specific identification is largely on the basis of probability,
since D. kallerti is the only species of the genus known from this
area, and the few characters exhibited agree. I have no doubt that
it is correctly identified with that species.

Pseudoeurycea bellii (Baird)

Oedipus platydactylus Baird, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 1, 1849,
p. 286 (nee Salamandra platydactylus Cuvier). (Type locality, Mexico.)

Spelerpes bellii Gray, Catalogue of the Batrachia gradientia of the British Museum,
1850, p. 46.

Pseudoeurycea bellii Taylor, Univ. Kansas Sci. Bull., vol. 30, 1944, p. 209.

A partly digested specimen of this species was taken from the
stomach of Thamnophis scalaris scaliger (Jan). The characteristic
red color markings of head and body identify this species without
question. The snake is no. 293 from the Xilitla region and is men-
tioned elsewhere in the paper under its name. This is a new record
of P. bellii for the state and the most northerly record on the eastern
part of the plateau.

Bolitoglossa platydactyla (Gray)

Salamandra platydactylus Gray, Supplement, in Griffith and Pidgeon's Cuvier's, The Ani-
mal Kingdom, vol. 9, 1931, p. 107. (Type locality, Mexico.)

Bolitoglossa platydactyla Taylor, Univ. Kansas Sci. Bull., vol. 30, 1944, p. 219.

The single specimen in the collection (no. 525) comes from the
Xilitla region. It displays all the usual characteristics of this spe-
cies. The dorsal coloration is uniform brownish-faun, with an elon-
gate, V-shaped, dark mark on the head and neck, followed by four
or five irregularly-placed dark flecks.

I suspect that this specimen came from a much lower elevation
than that of the town of Xilitla (2,200 ft.), since the species is not
known to occur much above 2,000 feet elevation.

Chiropterotriton sp.

A salamander taken from the stomach of no. 285, a young Tham-
nophus phenax phenax Cope, has the anterior part of the body more
or less intact, save that the tips of the toes have the terminal pads
missing, leaving the terminal bones exposed.

The following characters are evident: maxillary teeth 30-30, sub-

174 The University Science Bulletin

equal in size; premaxillary teeth, 8; vomerine teeth, 7-7, the series
beginning on the inner edge of the choanae; separated from each
other posteriorly by a distance equal to the narrowest diameter of
choanae, and from the palatine series by a distance equal to two-
thirds of the distance between choanae. Mandibular teeth 34-34.

First finger completely involved in web; others with small web
at base, leaving 2i/^ phalanges free. The costal grooves are 13.
The median part of the venter is without pigment; some pigment is
present on the chin.

Most of the other external characters are obscured by the process
of digestion, which makes specific identification uncertain. The
teeth are somewhat more numerous than in Chiropterotriton multi-
dentata Taylor. However, it is with this species that the form must
be compared. Not impossibly it represents an undescribed form.

SALIENTIA

Bufo horribilis Wiegmann

Bufo horribilis Wiegmann, Isis von Oken, vol. 26, 1833, pp. 654-655. (Type locality,
Misantla and Veracruz, Veracruz, Mexico.)

A young specimen (no. 527) was taken by Charles R. Shaw 3
miles north of Pujal, San Luis Potosi.

Bufo valliceps Wiegmann

Bufo valliceps Wiegmann, Isis von Oken, vol. 26, 1833, pp. 657-659. (Type locality,
Mexico.)

A series of 33 specimens was collected (nos. 439-471), chiefly in
April, May and June of 1947, although two specimens were found
as early as March 5 of the same year.

Most of the specimens conform to the typical characters and color
patterns of the species. Three, however (nos. 441, 456, and 459),
large gravid females, are of a very dark blackish color, lacking al-
most all trace of the typical color pattern of the other specimens.

Melanism has not been observed in other populations of this wide-
spread species.

SytThophus sp.

Two specimens of this genus are in the collection but unfor-
tunately both are in poor condition. One, no. 533, "found drowned in
a thunder- jug" is bloated and surface tissues are somewhat softened
and rotted. Consequently, some of the features, such as color
markings and skin characters, can be only doubtfully interpreted.
The second (no. 534), completely dried and shrivelled before being
placed in alcohol, likewise is difficult to identify.

Taylor: Herpetology of San Luis Potosi, Mexico 175

The species belongs in the section of the genus in which a tiny-
outer palmar tubercle is present, and the terminal pads on outer
fingers are distinctly wider than the digits. The choanae are small,
the diameter of one contained in the distance between them, five to
six times. One cannot be certain whether a web vestige is present
or not or whether the skin of the venter is areolate. The diameter
of the tympanum is about two thirds the diameter of the eye and
the eyelid is granular. The identity of the species is in question.
For one reason or another the following species of Syrrhophus in this
section can be eliminated: latodactylus, verruculatus, campi, gutti-
lahts, smithi, and leprus. If the specimens belong to a known species,
they are referrable to cystingnathoides or verrucipes with the
greater probability to verrucipes.

Three species are already laiown from the state. These are Syr-
rhophus guttilatus, S. latodactylus and S. campi. S. cystignathoides
has been reported with a question.

Eleutherodactylus hidalgoensis Taylor

Eleutherodactylus hidalgoensis Taylor, Univ. Kansas Sci. Bull., vol. 28, pt. 2, 1942, pp.
299-301, pi. 25, figs. 5-8; pi. 27, fig. 10.

This rare species has been known previously from two small male
specimens from near Tianguistengo, Hidalgo. Fortunately the
collection contains three specimens, two large females and one male,
all taken in a cave near Xilitla by Charles Shaw.

These specimens differ from the types in only small details. In
the two larger females, the head while proportionally the same,
shows no indefinite occipital fold, and the tympani of the females
are somewhat larger — more than half of the diameter of the eye. The
tympanum in the male is practically as in the type. On the tarsus
the inner fold is distinct and continuous in the large females, indis-
tinct in the male, and the tubercles on the outer side are faintly in-
dicated in the females and are not discernible in the male.

The color (in formalin transferred to alcohol) of no. 523 is gray
with some darker gray flecks. The tympanum is brown with a
lighter center. The snout and head are darker gray, with a faint
light interorbital line and two indistinct lighter diagonal lines or
bars behind eyes tending to inclose a darker triangle. The barring
on limbs is indistinct, the color dark gray. No. 524 is lavender to
purplish brown. The head pattern is discernible but indistinct, and
the back is clouded with darker. The bars on the limbs are more
distinct than in the type. In the male (no. 525) the w-shaped pat-
tern of the type is more plainly visible and the head markings are

176 The University Science Bulletin

likewise very distinct. The back is clouded or flecked with brown.
All of these three specimens are light flesh on the ventral surfaces
but there is a peppering of pigment visible under the lens.

The finding of these specimens in a cave is of great interest. It is
possible that crevices in rocks and caves, rather than trees, are the
normal habitat of the species and may account for the fact that only
a few specimens have been found.

Smilisca baudinii baudinii Dumeril and Bibron

Hy!a baudinii Dumeril and Bibron, Erpetologie generale ou histoire naturelie complete des
reptiles, vol. 8, 1841, pp. 564-5. (Type locality, Mexico.)

Hyla baudinii baudinii Stejneger and Barbour, Checklist of North American Reptiles and
Amphibia, 2d ed., 1923, p. 29.

Smilisca daulinia Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, 1865, Oct., p. 65;
footnote (^Hyla baudinii).

A series of 19 specimens (nos. 505-522, 532) was collected in May
and June of 1947. The earliest date is May 9, and the latest June
25. Many of these are gravid females. All were taken in the
Xilitla region or (one) "on the road to Xilitla."

Acrodytes syilomrna Cope

Hyla spilomma Cope, Proc. American Phil. Soc, vol. 17, July 20, 1887, p. 86. (Type
locality, Cosomoloapam, Veracruz, Mexico.)

Acrodytes spilomma Smith and Taylor, Univ. Kansas Sci. Bull., vol. 30, 1944, p. 64.

A single small specimen (no. 531) belonging to this species was
obtained at Pujal, November 21, 1946, by Charles R. Shaw. The
dorsal coloration is brown with rather broad cream lines beginning
behind the eye and extending back dorsolaterally about two-thu'ds
the length of the body. The limbs are lighter than the body, with
black flecks on the numerous pustules. It was found on a cornstalk.

Specimens have previously been reported from Rio Coy (near
Pujal) and Tamazunchale, in San Luis Potosi.

Hyla miotympanum Cope

Hyla miotympanum Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18, Mar., 1863, p. 47.
(Type locality, near Jalapa and Mirador, Veracruz, Mexico.)

A gravid female (no. 529), taken at Xilitla, is in the collection.
It differs from more typical specimens in having the tympanum
nearly half the diameter of the eye, while the typical form usually
has the tympanum only one third to one fourth the eye diameter.

There is a slight axillary web, and the first finger is opposed to
the other three. A continuous inner tarsal fold is present.

Specimens have previously been reported from Tamuzunchale and
Valles.

Taylor: Herpetology of San Luis PoTOsi, Mexico 177
Hyla arenicolor Cope

Hyla affinis Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, Apr., 1854, p. 61 (nee
Hyla affinis Spix, 1854). (Type locality, northern Sonera, Mexico.)

Hyla arenicolor Cope, , Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 6, July, 1866,
p. 64 (substitute name for the preceding species).

A single specimen (no. 528) was obtained by Robert Newman at
the San Luis Potosi Reservoir, March 28, 1947.

The specimen is ash gray with darker blotches. Specimens have
been previously reported from the state.

Rana pipiens Schreber

Rana pipiens Schreber, Der Naturforscher, Halle, vol. 18, 1872, p. 185, pi. 4. (Type
locality. Raccoon, Gloucester County, New Jersey, U. S. A.)

This widespread species is represented in Mexico by several forms
which may well be worthy of subspecific recognition. However, no
attempt has been made to allocate subspecifically a juvenile speci-
men in the collection (no. 530) taken at Villa del Reyes, San Luis
Potosi.

SAURIA

Anolis sericeus Hallowell

Anolis sericeus, Hallowell Proc. Acad. Nat. Sci. Philadelphia, 1856, pp. 227-228. (El
Encero de Jalapa, Veracruz.)

Seven specimens of this small anole are in the collection (nos. 561-
567). Three of these have the two orbital series separated by an
intercalated series of small scales. Four specimens have them sep-
arated save for one scale, which is in contact with its fellow. Four
are males with an ample gular appendage bearing a dark red brown
spot (said to be blue in life). The median portion of the appendage
is whitish.

This is a new record for the state.

Anolis petersii Bocourt

Anolis petersii Bocourt, Mission Scientifique aux Mexique el dans TAmerique Centrale;
Etudes sur les Reptiles et les Batraciens, Livr. 2, 1873, pp. 79-80, pi. 13, fig. 2; pi. 15,
figs. 11, 11a. (Type locality "la haute Vera Paz [Guatemala]"); Boulenger, Catalogue of
the Lizards in the British Museum, 2d ed., vol. 2, 1885, pp. 66-67 (Mexico).

Anolis biporcatus (non Wiegmann) Cope, Proc. Acad. Nat. Sci. Philadelphia, 1871, p. 215
(Mexico).

A single specimen, no. 560, was captured in the Xilitla region.
This form is rare in collections probably due to its habitat in forest
trees. It is the largest Mexican species of the genus Anolis.

The specimen, a young adult, measures 93 mm. snout-to-vent.
The type specimen is considerably larger, measuring 118 mm. This
is the first record for the state.
12—3659

178 The University Science Bulletin

Corythophanes hernandesii (Gray)

Chamaeleopsis Htrnandezii Gray, Synops. Rept. in Griffith's Cuvier's Animal Kingdom,
vol. 9, 1831, p. 45.

Corythophanes hertiamlesii Boulenger, Catalogue of the Lizards in the British Museum,
2d ed., vol. 2, 1885, p. 103.

A splendid series (nos. 407-438) of this rare casque-headed lizard
with a horizontal spine above the ear, was obtained by Mrs. New-
man in the Xilitla region. Two specimens, nos. 413 and 414, are
from the specific locality, Rancho Apetsco.

The specimens agree in structural detail with Boulenger's decrip-
tion {loc. cit.). However, there is variation in coloration between
the two sexes. Females have the sides of the snout light while the
eye is involved in a broad diagonal black line that reaches to the
edge of the tympanum. Three lines begin from lower edge of the
black stripe, and pass down across the jaws and then are deflected
backwards to breast. The bellies are usually banded with lighter
and darker brown, the latter may be broken into discrete spots.
Sides of head casque dark. One dark band on shoulder and four
more or less distinct short dorsal bands. The one on the rump is
more conspicuous than the others.

Males often have the black diagonal stripe involving eye, bor-
dered below with white. The sides of snout and entire lower jaw,
throat, and to a lesser extent the abdomen, light. A small protrud-
ing throat fan evident in males (to a somewhat lesser extent in fe-
males) . The sides of the casque are reddish brown with some black
reticulation.

Laemanctus serratus Cope

Laemanctus longipes (non Wiegmann) Dumeril, Arch. Mus., vol. 8, 1856, p. 512, pi. 21,
fig. 4; Bocourt, Arch. Mus., vol. 8, p. 114, pi. 17, fig. 4.

Laemanctus serratus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1864, p. 176. (Type
locality, not stated, presumably Mexico) ; Boulenger, Bull. Soc. Zool. France, 1877, p. 463,
pi. 7, fig. 3; Catalogue of the Lizards of the British Museum, 2d ed., vol. 2, 1885, pp.
104-105.

Three splendid specimens of this bizarre species are in the collec-
tion (nos. 337-339) taken at Xilitla by Marcella Newman. This
species has remained exceedingly rare in collections. In the exten-
sive collections made by Dr. Hobart M. Smith and me over a period
of years not a single specimen was taken. It is not known whether
the species is actually rare or only seemingly so. Its tree habitat
and green coloration in life would prevent easy observation and ac-
quisition by collectors.

Between the semicircular orbital series and the superciliary edge
there are about six rows of smaller unequal scales. The parietal

Taylor: Herpetology of San Luis Potosi, Mexico 179

area is constricted somewhat behind the orbits, and the lateral ser-
rate edges of the casque come together posteriorly at an angle. The
scales of the serrate edge are striate while all other scales are
strongly rugose or tubercular. A series of regular plates on the
snout between the two series of canthal scales extend back to the
level of the eyes. Despite the fact that the nasal is moved back and
separated by one or two scales from the rostral, the two most an-
terior of these may be interpreted as a pair of internasals in contact
mesially with each other, with the rostral anteriorly and with a
median azygos frontonasal posteriorly. The latter scale is bounded
laterally by two supranasal scales, the most anterior of which is in
contact with the internasals. The frontonasal is followed by a pair
of prefrontals. Although the supraorbital areas are not elevated,
each is bounded by a curved series of eight somewhat enlarged
scales, the two series normally separated by a single series of scales
(one specimen has two immediately following the prefrontals).

The limbs are extremely long; the adpressed hind limb reaches
beyond the tip of the snout. The subdigital lamellae have rounded
blackish, horny knobs, usually in a single series but under the joints
there may be one or more pairs. Two elongate keels appear on the
enlarged scales following the claws.

In life the specimens were nearly uniform green, save for the
white stripe from eye to shoulder, bordered above by a broad dark
line which narrows posteriorly. A series of brown spots are present
on the sutures between the labials, and a white stripe runs from
axilla to groin on the ventrolateral line. The tail is green but the
shade is a rather grayish green.

In two of the preserved specimens the green color is completely
gone, and is replaced by a reddish brown on the back and sides, on
which is superimposed five irregular diagonal blotches, each joining
its fellow on the back, the spots outlined in black or brown color.
The chin is flecked with red brown. The tail is indefinitely banded
with darker. The labials have a bluish cast, while the chin and ven-
ter are reddish purple with some greenish blue evident midventrally,
and posterior to the anus. A cream-white line runs from under the
eye to the shoulder and a cream spot or line is present on the side of
the neck, both of which are parallelled by a darker stripe. The ven-
trolateral white stripe (partly discontinued) is present on the ven-
trolateral surface.

The third specimen which has undergone a different preservation
in a heavy salt brine has retained most of its life colors.

180 The University Science Bulletin

Ctenosaura acanthura (Shaw)

Lacerta acanthura Shaw, General Zoology, vol. 3, pi. 1, 1802, p. 216. (Type locality
[restricted to] Tampico, Tamaulipas, Mexico.)

Ctenosaura acanthura Sumichrast, Arch. Sci. Phys. Nat., vol. 10, pp. 49-50; Bailey, Proc.
U. S. Nat. Mus., vol. 73, art. 12, 1928, pp. 9-16, pis. 3, 4.

Two specimens, a young (no. 344), and an adult (no. 345), are in
the collection from Ebano, San Luis Potosi.

The male is very dark, and little trace of the dorsal pattern re-
mains. The dorsal crest is from 8 to 13 mm. high, extending from
the back of the occiput to a point slightly back of the level of the
thigh. The spiny whorls of the tail are separated by three smooth
rows, at base (for 2 or 3 whorls) and by two for nearly half the
length. Beyond this point there is little differentiation in the trans-
verse rows. The femoral pores are 7-6.

Crotaphytus coilaris baileyi (Stejneger)

Crotaphytus baileyi Stejneger, N. American Fauna, no. 3, 1890, p. 103, pi. 12, fig. 1.
(Type locality. Painted Desert, Arizona.)

One specimen, no. 336, is from Presa de Gaudelupe, S. L. P.

The species of Crotaphytus occurring in Eastern San Luis Potosi
appears to agree more closely with the southwestern C. coilaris
baileyi rather than with C. coilaris coilaris which is the expected
eastern species. However, it differs from C. baileyi in the following
characters: The median scales between the supraocular areas are in
two enlarged series which are almost completely separated by an
intercalated series of scales. Only two pairs of scales touch for a
part of their length ; normally three or four pairs are in contact for
their entire length.

The head is grayish or clay color with small black dots scattered
on the top and sides. A double black collar is present, the anterior
part formed by a series of dots. The posterior part is continuous on
each side, but above is interrupted mesially. About six dim, narrow,
light, transverse bars are present on the back, the intervening areas
being dotted with rounded dim, gray spots. Between the shoulders
there is a transverse row of contiguous black dots; two other rows
across the back, the black dots in these, however, are separate and
not especially distinct. The chin is clouded or mottled with gray
and dull cream; the underside of the abdomen, and the underside of
the hands and feet are a uniform creamy white. The tail is bluish
gray with numerous small rounded darker spots.

Should the character of the intercalated median scale row on the
head prove constant in a series, this lizard might well be recognized
as a new form.

Taylor: Herpetology of San Luis Potosi, Mexico 181
Holbrookia maculata approximans Baird

Holbrookia approximans Baird, Proc. Acad. Nat. Sci. Philadelphia, 1858, p. 253. (Type
locality, "Lower Rio Grande.")

Holbrookia viaculata approximans Stejneger, N. American Fauna, no. 3, 1890, p. 109.

A specimen, no. 340, was captured five miles NE of Candido
Navarro, and no. 574 near the city of San Luis Potosi.

The specimens appear to be; typical and extend the known range
of the species from Central Chihuahua and Coahuila considerably
farther to the southeast.

Sceloporus jarrovii immucronatus Smith

Sceloporus jarrovii immucronatus Smith, Copeia, 1936, pp. 223-227. (Type locality "Ten
miles north of El Pinalito, Hidalgo, Mexico.)

Six specimens of this form (nos. 548-533) were taken in the gen-
eral area of Pendencia and Ciudad Maiz in eastern San Luis Potosi
(the elevation of no. 548 is 4,300 ft., two miles south of the village
of Pendencia.) The specimens, while agreeing in most characters, do
differ to some extent.

These constitute the first record of this subspecies in the state,
although Sceloporus jarrovii minor is known to occur in the west-
ern part. This latter form is characterized by a wide black nuchal
collar, six to eight scales broad. In this series the collars are only
three or four scales broad. The head and neck of the females have
numerous bluish white spots.

Sceloporus serrifer plioporus Smith

Sceloporus serrifer plioporus Smith, Field Mus. Nat. Hist. Zool. sen, vol. 26, July 27,
1939, pp. 212-214. (Type locality, Encero, Veracruz.)

A specimen from Ebano, no. 547, is referred to this form despite
certain differences from the type. It is a male and somewhat dis-
colored by formalin. The femoral pores are 10-11, and there are 32
scales from occiput to above vent. The posterior frontal is divided
longitudinally and separated from the interparietal.

The nuchal collar is about four scales wide and bordered behind
by a narrow light collar. A few lighter marks are present on head
and neck; the dorsal scales are normally mucronate. The tail has
about seven bluish bands separated by darker bands. The field
notes state: "back pale green and yellow; blue on sides of belly."

While this is the first record for the state, it is known from south-
ern Tamaulipas and northern Veracruz.

182 The University Science Bulletin

Sceloporus spinosus spinosus Wiegmann

Sceloporus spinosus Wiegmann, Isis, vol. 21, 1828, p. 370.

Sceloporus spinosus spinosus Martin del Campo, Anales Inst. Biol., Mexico, vol. 8, 1937.
p. 262; Smith, Field Mus. Nat. Hist., Zool. Ser., vol. 26, 1939, pp. 87-94, fig. 10.

One female specimen, no. 554, taken in the Charcas Region, in an
arroyo near Charcas Mine, 6,900 ft. elevation, is referred to this
species.

Sceloporus olivaceus Smith

Sceloporus olivaceus Smith, Trans. Kansas Acad. Sci., vol. 37, 1934, pp. 263, 277-279;
and Smith, Field Mus. Nat. Hist., Zool. Ser., vol. 26, 1939, pp. 110-118, pi. 13.

A large male, no. 555, from near Pendencia is referred to this spe-
cies. There are five supraoculars and the prefrontals are separated ;
the large bluish marks are separated mesially by an elongate light
stripe divided by a very thin black line ; the bluish spots themselves
have some indefinite dark markings; and the dorsolateral light
stripes are more or less distinct.

The scales of the head are black with narrow, greenish, transverse
markings. One line crosses the prefrontals, one crosses the supra-
oculars and another discontinuous line crosses the parietals. The
superciliary area has six or seven small greenish spots. Three defi-
nite greenish lines run from the eye down across the labials.

Sceloporus microlepidotus disparilis (Stejneger)

Sceloporus disparilis Stejneger, Proc. Biol. Soc. Washington, vol. 29, 1916, pp. 227-230.
(Type locality, Lomita Ranch 6 mi. N, Hidalgo, Texas.)

Sceloporus microlepidotus disparilis Dunn, Proc. Acad. Nat. Sci. Philadelphia, vol. 88,
1936, p. 472.

The single specimen (no. 342) from Xilitla is a male having the
femoral pores, 16-16; 68 dorsal scale rows from the parietal to the
posterior line of the thighs; and 67 scale rows around the middle of
the body.

The species has been reported previously from several localities
in the state.

Sceloporus variabilis variabilis Wiegmann

Sceloporus variabilis Wiegmann, Herpetologia Mexicana, 1834, p. 51. (Type locality
Mexico, by inference.)

Sceloporus variabilis variabilis Smith, Proc. Biol. Soc. Washington, vol. 47, pp. 127-129.

This is one of the most common lizards of the northern edge of the
plateau, and is represented by a large series of specimens (nos. 346-
406 and 568-576). Of these 14 are from Ebano, 1 from 21/2 miles
south of Pendencia, while the remainder are from the Xilitla region.
The species appears to remain active throughout the winter. In

Taylor: Herpetology of San Luis Potosi, Mexico 183

November, 14 were taken; December, 1; January, 5; February, 3;

March, 2; April, 28; and May, 18.

Specimens from Ebano and Pendencia have the femoral pores
small, and the rim of the pore scale is broken posteriorly. Those
from Xilitla have the pores twice as large as the Ebano specimens
and the rim of the scale surrounding the pore is usually unbroken.

Cnemidophorus gularis gularis (Baird and Girard)

Cnemidophorus gidaris Baird and Girard, Proc. Acad. Nat. Sci. Philadelphia, 1852, p.
128 ("Indianola and the Valley of the Rio Grande del Norte").

Cnemidophorus gularis gidaris Cope, Proc. Acad. Nat. Sci. Philadelphia, 1892, p. 334.

The subspecies Cnemiidophorus gularis gularis is represented by
six specimens (nos. 535-540) as follows: four from Ebano, one from
the "Charcas Region, desert at Morelos Mine," and one from the
"San Luis Potosi Reservoir."

The younger specimens are typically striped. In the largest, and
perhaps oldest, the stripes are missing and are replaced by numerous
blue-white punctations scattered on the dorsal surface.

Aineiva undulata podarga Smith

Ameiva undulata podarga Smith, Univ. Kansas Sci. Bull., vol. 31, pt. 1, 1946 pp. 40-43,
figs. ID, 2A.

Six specimens (nos. 541-546) were obtained in the Xilitla region.
The specimens appear to be typical of this form.

Xenomurus newmanorum sp. nov.

Type: No. 499 Louisiana State University. Collected in the
Xilitla region, San Luis Potosi, Mexico.

Paratypes: Nos. 489-498; 500-504, L. S. U., same locality.

Diagnosis: Related to Xenosaurus grandis but with the following
differences: supraocular scales smaller, never forming a continuous
series of three or four, that are much wider than long; tail a little
shorter and slenderer with 2-4 scales fewer in a whorl about tail
near base; arm above nearly uniformly covered with large rounded
scales, somewhat conical on dorsal surface, rather than with the
tubercles spaced with the distance between nearly equal to diameter
of the base of the tubercle ; a pair of light spots beginning between
auricular membranes and extending back for a distance less than, or
equal to, their width ; a broad distinct stripe begins near tympanum
that runs up and back, approaching but not meeting its fellow be-
hind posterior level of arm insertion ; venter nearly uniform grayish
white, instead of cream with numerous quadrangular black spots.

184 The University Science Bulletin

Description of the type: Dorsal outline of the head triangular
or arrow shaped, the temporal region slightly curving and bulging
out in front of the tympanum, which faces outward and backward;
the canthus rostrailis is poorly developed and there is no canthus
temporalis; rostral narrower than mental, ll^ times as wide as high,
highest at median point, notched laterally above; no distinctly en-
larged medial scale touching rostral ; circumorbital rings moderately
developed, separated by a single series of roughened scales; scales
of dorsal surface of head rough, keeled, ridged, or somewhat con-
ical, those in intemasal and temporal regions largest and with a
few scattered minute granular scales between; in temporal region,
scales distinctly conical, separated by irregular rings of minute
granular scales; median supraocular scales somewhat enlarged but
not forming regular series, often subcircular and moundlike; super-
ciliaries 11-11; 13-13 suboculars; nostril in single nasal; supra-
labials 13-13; infralabials, 11-12; a series of scales bordering su-
pralabials above heavily ridged posterior to nasal; a row of en-
larged scales begin behind eye and run back on temporal region,
usually bordered by one row of scales on each side also somewhat
enlarged.

Body covered with rounded or oval somewhat conical scales sep-
arated by smaller similar scales mixed with minute granules of
varying size.

A skinfold on neck, with often another indistinct one preceding
it; an average count of about 43 scales in a line between neck fold
and tip of chin (counts may vary three or four scales if made at
different points) ; beginning at about level of axilla, a vseries of trans-
verse scale rows cross the ventral surface^ terminating posteriorly
at about the level of the groin. These individual scales are quad-
rangular and there are 18-20 in the widest rows; on sides they
merge into the more rounded lateral scales; a lateral skinfold from
axilla to groin along sides, running close to ends of ventral series.

On median ventral line 78 scales from base of tail to tip; 107
scales on dorsal part of tail; tail surrounded by whorls of scales,
every other whorl breaking up into two rows on the sides so that
the dorsal count is usually nearly a third greater than ventral;
ventral part of tail compressed and narrowed.

Dorsal surface of arm regularly scaled, the tubercles in contact,
with occasional granules in the interstices; on the posterodorsal
part of thigh, tubercles are farther apart, not in contact.

Tympanum smooth not covered with granules, higher than wide,
its greatest diameter equal or very slightly less than length of eye

Taylor: Herpetology of San Luis Potosi, Mexico 185

orbit; fingers and toes well developed, somewhat angular, claws
strong; 32 scales under fourth toe; toes of hind limb reaching half
way between elbow and arm insertion when limbs are adpressed to
body; maxillary-premaxillary teeth, 23 on each side.

Color: Head with a ground color of brownish gray; numerous
dark brown flecks forming indistinct lines running back from eye
and a large more or less distinct dark mark in the occipital region;
back with five or six broad brown bands with deep black irregular
edges and lineated series of black tubercles ; eight dark bands round
tail, the under parts of bands split with lighter color; the brown
bands are separated by bluish gray bands that are diagonal on the
sides and are interrupted medially on back by a very narrow streak
of brown; these light bands may be edged with black and there
may be some scattered black tubercles; chin, throat and venter
dirty white or slightly bluish white. (Under a lens there is to be
seen a fine, nearly uniform, scattering of pigment.)

The brown neck band is somewhat triangular; the middle one on
the back tends to be partially divided by an extra gray band. Arms
and legs indefinitely flecked with brown, black, and gray above,
nearly uniform gray below; digits indefinitely banded with gray.

Measurements in nun.: Snout to vent, 112; tail, 106; head width,
25; head length to front edge of tympanum, 31; axilla to groin, 51;
snout to forearm, 44; body width, 23; arm, 40; leg, 50,

Remarks: The differences between this form and the typical one
occurring in Veracruz near Cordoba are numerous but not of great
magnitude. Certain of these have been pointed out in the diag-
nosis; the following differences exist between Xenosaurus grandis
and newmanonim: In grandis the head is more distinctly trian-
gular, the canthus temporalis more distinct, and the sides of the
temporal region more nearly vertical; the tympanum is entirely
covered with a thin scaly membrane (apparently absent in new-
manorum or if present, lacking scales) ; the area of small scales
behind the vent is larger, arranged in seven or eight rows (usually
only four, and the scales back of anal lip larger in newmanorum) ;
the preanal scales more uniformly developed. In the basal portion
of the tail every fifth whorl is divided, but in the remainder of the
tail every other row splits. This takes place lower down on the
side, often only the ventral median pair being undivided. The lower
labials have a low lateral ridge, less conspicuous than that in new-
manorum. The enlarged row of ridged scales below the infralabials

186 The University Science Bulletin

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Taylor: Herpetology of San Luis Potosi, Mexico 187

are wider than in nemmanorum. The first and second pairs of chin-
shields are in contact with the infralabials, while those following
are separated from the labials by the series of ridged scales. (In
newmanorum there are eleven specimens in which only first pair
touches labials; three in which one of second pair touches; in two,
all the scales of the chin are distorted and seemingly abnormal.)

The genus is known from Mexico and Guatemala. Xenosaurus
grandis occurs in the region about Cordoba, Cuautlapan, Huanusco,
Orizaba and Tehuantepec in Mexico; X. rackhami is a Guatemalan
species.

The specimens of X. grandis that I collected at Cuautlapan were
ensconced between lamina of rock, where softer layers had been
weathered away leaving narrow spaces between harder layers; it
is not known whether a similar habitat is occupied by X. new-
manorurn.

The species is named for Mr. Robert Newman and Mrs. Marcella
Newman who obtained the specimens.

Barisia imhricatus ciliaris Smith

Gerrhonotus levicollis ciliaris Smith, Proc. U. S. Nat. Mus., vol. 92, 1942, pp. 3C5-367.
(Type locality. Sierra Guadalupe, Coahuila, Mexico.)

Barisia imhricatus ciliaris Tihen, Amer. Mid. Nat., vol. (In press.)

This form, previously known from the "mountains near San Luis
Potosi [city]" is represented in the collection by a specimen (no.
341) taken at an elevation of 7,000 feet between Llano de Coneja
and Llano de Garzas in the Cerro Coneja region. It is a gravid
female containing 11 well-developed embiyos, probably about ready
for birth since the yolk material seems to be entirely absorbed.
One embryo measured 36 mm. from snout to vent, the total length
being 98 mm.

The following characters are diagnostic:

Dorsal scale rows 16, the six median being strongly keeled, the
adjoining rows lightly keeled; 12 ventral scale rows; a single pair
of chinshields in contact medially, the upper largest. The meas-
urements (in mm.) are: total length, 293; snout to vent, 115; vent
to tip of tail, 178. The adpressed limbs fail to touch.

The azygos postmental present on the specimen is perhaps ab-
normal for this form since only in four of the eleven young is it
present; and while two loreals are present in this specimen, one is
the more usual number.

188 The University Science Bulletin

Gerrhonotus liocephalus lowcryi Tihen

Gerrhonotus liocephalits loweryi Tihen, Trans. Kansas Acad. Sci., vol. 51, 1948, pp. 302-305.

This spendid series has recently been studied and named by Dr.
Joseph Tihen. The type is LSU no. 480. The paratype series,
LSU nos. 472-479, 481-485, 485a, 486-487. All are from the Xititla
region. One specimen from Ciiidad Maiz, no. 488, has been ten-
tatively referred to this form, but it differs in a number of points.
It is possible that these presumed differences are juvenile char-
acters.

Eumeces tetragrammus (Baird)

Plestiodon tetragrammus Baird, Proc. Acad. Nat. Sci. Philadelphia, 1858, p. 256. (Type
locality, Lower Rio Grande River.)

Eumeces tertagrammus Cope, Bull. U. S. Nat. Mus., no. 1, 1875, p. 45; Taylor, Univ.
Kansas Sci. Bull, vol. 23, 1935 (Aug. 15, 1936), pp. 298-304, fig. 46.

An example of this species no. 343 collected at Ebano appears
to be typical in coloration, and the scale data fall within the known
limits of variation.

TESTUDINATA

Terrapene mexicana (Gray)

Cistuda {Onychotria) mexicana Gray, Proc. Zool. Soc. London, 1848 (Feb., 1849), pp.
16-17, pi. 2. (Type locality, Mexico.)

Terrapene goldmani Stejneger, Proc. Biol. Soc. Washington, vol. 46, 1943, pp. 119-120.
Terrapene mexicana Miiller, Zool. Anz., Bd. 113, Heft 5/6, 1936, pp. 97-114, figs. 1-4.

Three specimens of Terrapene are in the collections; no. 332
Valles, S. L. P. ; nos. 333 and 334 being from Ebano, S. L. P. ; while
no. 335 was taken at Pujal, S. L. P.

Despite rather extraordinary differences in the general appear-
ance, all three are referred to Terrapene mexicana (Gray), a species
described nearly a hundred years ago from a specimen of unknown
provenance in Mexico. The type specimen has a dark brown cara-
pace, the dorsal and lateral plates being spotted and rayed with
cream. The scales on the arms are dark, and most of those on the
anterior face of the arm bear a yellow or orange spot. Four claws
are present on the fore foot. There are three claws on the hind foot.

Stejneger has described a Terrapene goldmani from "Chijol (or
Chijoles), southeastern comer of the State of San Luis Potosi,
Mexico; in the coastal plain." This species also has only three
claws on the hind foot. The carapace is nearly uniform "clay-
color" with well-defined broad dark-brown margins to each plate
along the seams. The color of the arms and legs is not mentioned.

Miiller {loc. cit.) has reviewed a series of Terrapene collected for

Taylor: Herpetology of San Luis Potosi, Mexico 189

the most part in the region about Tampico, Veracruz. He has con-
cluded that Terrapene goldmani and Terrapene mexicana are syn-
onyms. Whether he is correct in his interpretations cannot be de-
cided on the basis of these specimens. The more outstanding
characteristics of each of these is given.

No. 332 (collected by Chas. Shaw, Valles, San Luis Potosi, Oct.
11, 1946). This specimen has a high, laterally compressed cara-
pace, with a continuous, fiat crest on the second, third and fourth
vertebral scales. The keel can be distinguished also on the first
vertebral. A small nuchal is present. The marginal scales flare
out posteriorly and are strongly serrate. The foot has three claws.
The upper parts of the sides slope sharply, giving the carapace a
tectiform shape.

The general color is yellowish olive, the sutures of all the plates
being dark, black brown. The plastral plates are similarly col-
ored, in that the posterior edges of the scutes and the median su-
tures are dark brown. The chin and lips are dull brownish-white
without any markings. An indefinite grayish spot is present on
each side of the mandible. The neck, arms, legs and feet are uni-
form olive-brown without trace of yellow spotting on scales. The
top of the head is brown with some marbling of dirty cream. The
tip of the snout and maxilla are light olive. The area about the
nostrils is yellowish tan.

No. 333 from Ebano is the largest specimen. The carapace slopes
regularly on the sides. The sutures are bordered by wide areas of
dark black-brown. The scales on the front of the arms have yellow
spots, or are yellow with darker edges. The top of the head and
to a lesser extent the neck above, and side of head is covered with
numerous black spots. A pair of vertical lines is present on the
front of the beak which is very slightly notched. The plastron has
a broad dark area bordering the sutures, but some yellowish areas
are present near the outer border. The chin is whitish.

No. 334, from the same locality, lacks the dark stripes along the
sutures but does show what appear to be remnant spots and rays of
dark brown on a lighter olive ground color. The larger scales on
the arms are blackish with yellow centers. The top of the head is
more or less uniform bluish gray.

The specimen from Pujal (no. 335) has the dark stripes along
the sutures rather indistinct and some indistinct darker markings
suggesting spots or rays. There is a very small amount of dark
brown on the carapace, scattered in irregular areas along the su-
tures. The throat is strongly reticulated with black.

190 The University Science Bulletin

A very slight longitudinal convexity is discernible crossing the
upper half of the costals. The arms are spotted with yellow.

Measurements oj Terrapene mexicana (Gray)

no. 333 no. 33i no. 335 Jio. 332

Length of carapace 166 150 153 132

Length of plastron 164 138 142 126

Width of carapace 128 119 121 113

Width of plastron 94 85 87 81

Greatest height of carapace and plastron 88 72 78 82

SERPENTES

Leptotyphloqjs mycrpicus myopicus (Garman)

Stenostoma myopicum Garman, Mem. Mus. Comp. Zool., vol. 8, 1883, pp. 6, 130, 131.
(Type locality Savineto, near Tampico, Tamaulipas.)

Leptotyphlops myopicus ?nyopicus Smith, Field Mus. Nat. Hist. Publ., Zool. ser., vol.
29, 1944, p. 146.

An individual of this very rare species (no. 201) taken Novem-
ber 13, 1946, by Marcella Newman at Ebano, yields the following
data :

Total length, 214 mm.; tail, 10.5 mm., head width, 3.5 mm., body
width, 3.5 mm. Dorsal scales (transverse scale rows) total 120;
on tail 14, the first pair following anus divided ; others single. Anal
a large undivided scale; 14 scales in a row about body. Width of
rostral slightly more than one-third greatest width of head, ex-
tending back to level of eyes or slightly beyond; nasal completely
divided, the suture passing from nostril reaches first labial, the upper
part of scale much larger than lower, and separated from its fellow
of the opposite side by a small prefrontal; supraocular small, much
above level of eye; frontal about size of prefrontal; parietals wide,
the anterior reaching the last (third) labial, each separated from
its fellow of the opposite side; labial border formed by rostral,
nasal, two anterior labials, the ocular, and third labial; rostral oc-
cupying greatest space, the third labial a slightly shorter space on
the labial border.

The lower lip of this specimen is strongly notched mesially, with
four infralabials on each side of the diminutive mental. The fourth
labial is concealed entirely below the supralabials.

Constrictor constrictor imperator (Daudin)

Boa imperator Daudin, Histoire naturelle . . . Reptiles, vol. 5, 1803, pp. 150-152.
(Type locality, Mexico.)

Constrictor constrictor imperator Ihering, Rev. Mus. Paulista., vol. 8, 1910, p. 321.

Two specimens, nos. 202-203, are in the collection. The data on
these two specimens are, respectively: Ebano, Xilitla; female, fe-

Taylor: Herpetology of San Luis Potosi, Mexico 191

male; scale formula, 54-64-38; 56-66-36; ventrals, 230, 236; sub-
caudals, 53, 41+; supralabials, 21-19, 19-20; infralabials, 22-?,
23-22; total length, 1700, 1830; tail length, 160, 162+.

Adelphicos quadrivirgatus quadrivirgatus Jan

Adelphicos quadrivirgatus Jan, Arch. Zool., vol. 2, 1862, pp. 18-19, pi. 8; Elenco sist.
ofidi, 1863, p. 32; Jan and Sordelli, Iconographie Generale des Ofidiens, livr. 11, 1865,
pi. 3, fig. 5. (Type locality, Mexico.)

Adelphicos qiiadrivirgatus quadrivirgatus Smith, Proc. Rochester Acad. Sci., 1942, pp.
188-192, figs. 3 and 6.

Smith, loc. cit., notes that seven specimens of this species are
known, four of them having been taken in Mexico, three in Chiapas.
The known range extends from central Chiapas to Jicaltepec, cen-
tral Veracruz. Two mutilated specimens in the collection, no. 205
from Ebano, and 206 from Xilitla, trace the range from Jicaltepec
about 90 miles farther to the northwest. The latter half of the
body of the Ebano specimen is missing; consequently the scale
counts of this part are unknown, and the sex cannot be determined.

The general color is browTiish-faun, most of the scales having
more pigment on the edges than in the center. A nearly continuous
black lateral line runs between the second and third scale rows,
reaching to near the tip of the tail. A dorsolateral line begins near
the parietal, and follows a course through the middle of the fifth
scale row. The median dorsal scale row is pigmented slightly
more than adjoining rows, but would not be described as bearing
a stripe. The labials are cream, save for their brownish upper
edges. The venter is white save for a fine zigzag line under tail.
The top of the head is gray, the scales flecked with black.

The scale formula for no. 206 $ is 15-15-13; ventrals, 148; supra-
labials, 7-7; infralabials, 5 (the anterior chinshield bordering the
mouth also); no preoculars, postoculars, 2; temporals 1+1; one
loreal entering eye; anal divided. Tip of tail absent.

The mutilated specimen (no. 205) disagrees in no character that
can be discerned.

Ficimia streckeri Taylor

Ficimia streckeri Taylor, Copeia, 1931, p. 5-7. (Type locality, 3 miles east of Rio Grande
City, Texas); Mulaik and Mulaik, American Mid. Nat., vol. 29, no. 1, May, 1943, pp.
796-797.

Ficimia olivacea streckeri Smith, Publ. Field Mus. Nat. Hist., Zool. Ser., vol. 29, 1944,
p. 139; Smith and Taylor, Bull. U. S. Nat. Mus., no. 187, 1945, p. 63.

A series of 10 specimens of this burrowing species was acquired
in the Xilitla region. It is important since it gives a better idea of
the range of variation in the form.

192 The University Science Bulletin

The preoculars are uniformly 1-1; the supralabials 7-7 save in
no. 221, which has the first three fused into a single scale; infra-
labials, 7-7, save in one specimen with 7-8 on one side. There is

no

loreal.

Data

on Ficimia

streckeri

Taylor

Post-

Scale

Total

Tail

No.

Sex

ocular

formula

Ventral;

3 Caudals

length

length

207

$

1-1

19-17-17

146

34

139

22

208

$

2-2

19-17-17

144

36

418

69

209

S

2-2

17-17-17

148

36

424

66

210

s

2-2

17-17- ?

154

37

443

68

211

$

2-2

19-17-17

150

36

251

38.5

212

$

1-1

18-17-17

147

38

470

75

213

s

2-2

19-17-17

150

36

483

74

214

$

2-2

19-17-17

144

36

372

59.5

215

9

1-1

17-17-16

155

32

390

55

216

9

2-2

19-17-17

154

30

250

31.5

217

9

1-1

19-17-17

157

34

450

60

These specimens differ in some characters from those in the type
area in south Texas. Muliak and Muliak, loc. cit., give data on a
series of specimens (over 30?). A comparison of their ventral
counts and subcaudal counts with the Xilitla series is given. Num-
bers in parentheses are averages.

Texas

s $

128-144

35-40

• ■ • •

Xilitla

$ $

144-154

(148)

34-38

(36)

Texas

9 9

144-150

....

30-34

• • < •

Xilitla

9 9

154-157

(153.3)

30-34

(32)

In Texas specimens the supralabial number is 7; infralabials, 8
most frequently, with 7-7, or 7-8 less often; preoculars, 1; post-
oculars, 1 (one exception) ; temporal formula, 1+2.

The Xilitla specimens have, normally, supralabials 7 ; infralabials,
7 (one exception, on one side); preoculars, 1; postoculars, 2-2 in
7 and 1-1 in 4 cases; the usual temporal formula is 1+2.

Smith, loc. cit., has recently associated this form with Ficimia
olivacea as a subspecies on the basis of presumed intergrades be-
tween the two species. He is followed in this by Smith and Taylor,
loc. cit. However, this lot of material seems to show no evidence
of intergradation, and even with the loss of much of the dorsal color
pattern as occasionally occurs, there is no approach to the more
narrow, flatter-headed species with a differently colored, more
slender body.

Schmidtophis genus novum

Maxillary teeth eight or nine, decreasing in size posteriorly. Head
moderately elongate, scarcely distinct from neck. Eye small with
an elliptic pupil. Scales without striations, but with dim keels in
15 rows; large paired marginal "pits" on many if not all scales of

Taylor: Herpetology of San Luis Potosi, Mexico 193

anterior part of body; scale row formula, 15-15-15; ventrals
rounded; anal single; tail short with subcaudals in two rows; pre-
frontals fused entering eye; loreal entering eye, no preoculars; su-
praocular and postocular present, not fused; no anterior temporals.
Nasal at least partially divided.

The relationship of the genus is with Geophis and Chersodromus.

The genus is named for Karl Patterson Schmidt.

Schmidtophis rubriventris sp. nov.

Type: Louisiana State University No. R. 577. Collected near
Xilitla (Xilitla Region) San Luis Potosi, Mexico, by Charles Shaw.

Diagnosis: Characters of the genus. A very broad rostral one
third broader than high, one subtriangular supraocular; one post-
ocular much higher than long; frontal as long as its distance from
intemasals; body scales lightly keeled, save entire outer row, and
part of second row; no primary temporal. Body black, venter pink,
a white band across middle of parietals joining the white of throat.
Scales in 15 rows; 6-6 supralabials. Ventrals, 125, caudals, 41.

Description of type: Rostral broader than long, the part visible
above broad and narrow, its posterior border forming a very oblique
angle; internasals small, about one and one-half times as wide as
long; prefrontals fused together, the anterior border of scale some-
what convex, the posterior border somewhat concave; frontal rather
small, subtriangular, the anterior border slightly curved, its width
greater than its length, its length equal to its distance from inter-
nasals; supraoculars rather large, subtriangular; length of parietals
equal to their distance from intemasals; nasal large, at least par-
tially divided, the length of its rostral border greater than its dis-
tance from eye, the anterior part greater in area than posterior part;
loreal large, nearly rectangular (the lower edge slightly angular),
a little longer than high, entering eye; postocular at least twice as
high as long; fifth labial broadly in contact with parietal; no pri-
mary temporal, one secondary temporal shaped like a body scale;
six supralabials, in the following ascending order of size: 1, 2, 3, 4,
6, 5; the third and fourth supralabials enter eye; seven infralabials,
in following ascending order of size: 2, 1, 3, 4, 6, 7, 5, the first pair
in contact behind the small mental; first pair of chinshields rela-
tively large, bordered by five infralabials; second pair of chinshields
less than half size of first pair, in contact with each other. The
scales are smooth and lack any trace of striation; and the keel is
weak, but somewhat more pronounced posteriorly; outer row and

13—3659

194 The University Science Bulletin

part of second not keeled. Scale formula, 15-15-15; ventrals, 125;
anal undivided; subcaudals, 41-|-1.

Color: Color deep slate-black on dorsum and sides; a white band
with irregular borders crosses the back part of head on parietals
passing down on throat where it widens, covering about four ven-
trals and the posterior chinshields; head and supralabials black;
infralabials grayish black with a light area on mental and light
areas on last three infralabials; first pair of chinshields whitish
with grayish black anterior borders; ventrals and subcaudals uni-
form pink, the outer edge of ventrals colored like lateral scales.

Teeth: The short maxillary bears six teeth, with places for two
or three more. Anterior teeth larger, very slender, curved, growing
smaller posteriorly.

Measurements in mm.: Total length, 308; tail, 68; width of head,
6.3; length of head, 8; diameter of eye, 1.2.

Remarks: This novelty seems to mimic Chersodromus leibmannii
(Reinhardt), which may occur in the same locality but the nearest
point at which the latter has actually been found is in central Vera-
cruz about 180 miles to the southeast.

The chief generic characters by which it differs from Cherso-
dromus are: smooth instead of finely striated scales; absence of an
anterior temporal allowing the fifth supralabial to form a long con-
tact with the parietal; the smaller series of maxillary teeth, de-
creasing in size posteriorly; and the reduction of the scale rows
from 17 to 15 ; the supraocular and postoculars separate rather than
fused.

There is some similarity to certain members of the genus Geophis,
especially those lacking anterior temporals. However the fusion of
the prefrontals in this species would prevent confusion with these
forms.

Geophis multitorques (Cope)

Rhabdosoma multitorques Cope, Proc. Amer. Philos. Soc, vol. 22, 1885, pp. 384-385.
(Type locality, Zacualtipan, Hidalgo.)

Geophis multitorques Ciiinther, Biologia Centrali-Americana, Reptiles, 1893, p. 93; Smith,
Proc. New England Zool. Club, vol. 18, 1941, pp. 53-55.

Two specimens, nos. 218 (young) and 219 (adult female), from
the Xilitla region, are referred to this species. The scale data of
the two specimens agree for the most part.

The scale formula is 17-17-17; supralabials, 6; infralabials, 7; no
preocular; postocular, 1; temporals, 1+2; second pair of chinshields
in contact. The ventrals of no. 218 are 174, of no. 219, 177; caudals
are 30 in both. The measurements (in mm.) are, respectively:
total length, 171, tail, 21; 443, tail, 51.5.

Taylor: Herpetology of San Luis Potosi, Mexico 195

Above, the color is brownish black to black, the skin between the
scales being lighter. A parietal collar is present in no. 218 covering
about half the length of the parietals. It crosses the sides of the
mouth and widens to encompass the greater part of the underside
of the head. In both, the ventral surface of the body is black or
blackish brown with numerous white areas. These are sometimes
arranged as alternate spots, often contiguous or narrowly separated,
sometimes confluent. The undersurface of the tail is nearly uni-
form black.

Ninia diademata plorator Smith

Ninia diademata plorator Smith, Copeia, 1942, no. 3, Oct. 8, pp. 152-154. (Type lo-
cality, Durango, Hidalgo, Mexico.)

In defining this subspecies, Smith, loc. cit., had data from but two
specimens. The present series of six from the Xilitla region of San
Luis Potosi contribute to a better understanding of the variation
in the form.

The color of all the specimens agrees with that given for the type.
The scale formula, 19-19-19, is invariable (stated in error by Smith,
"18 scale rows" in the diagnosis, but giyen correctly in the descrip-
tion). The supralabials are 6-6 in five, 6-5 in one, specimen. Thus
the count of five supralabials in the type is apparently not the nor-
mal condition for the subspecies. Six infralabials are present. The
temporals are constantly 1 -(- 2, no preocular, 2 postoculars, the up-
per the larger. Four labials touch the anterior chinshields. In these
characters, agreement with the type is complete. There are invari-
ably two pairs of chinshields, the anterior much the larger.
Data on Ninia diademata plorator Smith

Total

Tail

No.

Sex

Locality

Ventral

Caudal

length

length

220

$

Xilitla region

133

83

254

82

221

5

Xilitla region

130

81

315

100

222

5

Xilitla region

129

74

290

88

223

9

Xilitla region

137

73

312

89

224

$

Xilitla region

125

88

302

98

225

$

Xilitla region

128

93

225

78

The variation in the ventral and caudal scale counts (including
data on the types) is as follows: 123-133, 83-93 for males; 129-137,
73-81 for females.

The counts in the form Ninia diademata diademata are distinctly

higher (136-145; 86-97 for males; 136-159; 79-96 in females). The

averages for these data are:

N. d. plorator 126.8—87.6 for males; 132—72 for females.

N. d. diademata 141.8—91.4 for males; 149.5—86.3 for females.

The stomachs of three of the specimens contained remains of
small slugs. One specimen contained oviductal eggs.
This is the first record of the species for the state.

196 The University Science Bulletin

Rhadinaea crassa Smith

Rhadinaea crassa Smith, Proc. Biol. Soc, Washington, vol. 55, 1942, pp. 190-191, fig.
3, 4. (Type locality Durango, Hidalgo, Mexico.)

This recently described species has been known previously from
five specimens, one of which, a paratype, is from Ciiidad Maiz, in
San Luis Potosi. Three more specimens, nos. 262, 263, and 264, have
been collected by Mrs. Marcella Newman in the Xilitla region. All
are males and appear to have a larger series of ventrals than the
more typical, described specimens.

The following characters are invariable in the three: supralabials,
8-8; infralabials, 9-9; preocular, 1; postoculars, 2; temporals, 1+2;
loreal, 1; anal, 2; five scales touch the anterior chinshields; scale
formula, 17-17-17.

Data on Rhadinaea crassa Smith

No.

Sex

Locality

Ventrals

Caudals

Total
length

Tail
length

262

$

Xilitla

185

91

587

176

263

$

Xilitla

181

94

545

162

264

$

Xilitla

184

92

587

176

The color patterns of these three specimens agree with each other,
and likewise with that of the type specimen.

Females of a snake species usually have a higher average number
of ventral scales than the male. In the three typical specimens of
R. crassa from which data on ventral counts are available, all are
females, and the range of the ventral counts is 166-173, average
170.6. It is presumed that typical males would have a lower
average.

In these Xilitla specimens the range for three males is 181-185
with an average of 183.3, a difference of at least 13 ventral scales
greater than the females of the species. It may also be larger than
the average for typical males.

Rhadinaea decorata (Giinther)

Coronella decorata Gunther, Catalogue of the snakes of the British Museum, 1858, pp.
35-36. (Type locality Mexico.)

Rhadinaea decorata Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 15, 1863, p. 101.

This species, not previously known from the state, is represented
by three male specimens (nos. 265-267), all from the Xilitla region.
They extend the known range approximately 100 miles to the north-
west. To the south the species reaches to Costa Rica.

These specimens agree with the more southern specimens in hav-
ing the scale formula, 17-17-17; supralabials, 8-8, infralabials, 10-
10, five touching the first chinshields; preoculars, 1-1, postoculars,
2-2, temporals, 1-f 2.

Taylor: Herpetology of San Luis Potosi, Mexico 197

Only one specimen, no. 266, has the tail complete. The ventral
counts for the specimens nos. 265, 266, and 267 are, 126, 118, 121,
respectively. The caudals for no. 266 are 121. The ventral color-
ation of the preserved specimens is creamy white. The color in life
was not recorded.

The stomachs of two specimens contained the partially digested
remains of earthworms.

Rhadinaea marcellae sp. nov.

Holotype: Louisiana State University no. 270, from Xilitla re-
gion, San Luis Potosi, Mexico, May 12, 1947. Collected by Mar-
cella Newman.

Diagnosis: Dorsolateral light line on head beginning on the
snout, running through upper edge of eye to behind mouth angle,
then back and upward to join the lateral body stripe, which oc-
cupies the fifth rows and is composed of a series of disconnected
light spots each covering about half of a scale; a nuchal white
black-edged collar connecting the ends of the lateral body stripes;
head with a somewhat elaborate stippled pattern; outer edge of
ventrals dark, the area intensified at the posterior outer edges of
the scales. Black dots on most of the supralabials, and smaller
spots on mental and three first infralabials; 2 preoculars, the lower
small; ventrals, 128; caudals, 78; four infralabials touch first chin-
shields.

Description of the type: Rostral wider than high, visible only
as a narrow line when seen from above; internasals more than half
of the length of the prefrontals; frontal one third longer than wide,
about one third longer than its distance from the tip of the snout;
parietals a little longer than their distance to the tip of the snout;
nasal divided; loreal quadrangular; two preoculars, the lower very
small, separating the third labial from the orbit; two postoculars,
the upper more than three times area of lower; anterior temporal
large, followed by two smaller temporals; 8-8 supralabials, the
sixth and seventh very large and subequal in area, the fourth and
fifth form the lower border of the orbit; eye very large, greater
than its distance to the anterior edge of the nostril ; 8-9 infralabials ;
two pairs of chinshields, the anterior broader, but somewhat shorter
than the posterior; 4 labials touching anterior chinshields. Scale
formula, 17-17-17. Ventrals, 128; caudals, 78; anal divided.

Total length, 290 mm., tail, 93.

Color and markings: A line from tip of snout through upper part
of the eye, then diagonally to behind mouth- angle, then back and

198 The University Science Bulletin

up joining the lateral body line; a black-edged white nuchal collar
joining the ends of the dorsolateral lines which are very narrow
(about the width of half a scale), formed of a series of small sep-
arate whitish spots each about half the area of one scale, and fol-
lowing the fifth scale row. Below this a veiy narrow dark line,
less than half a scale in width, and covering part of lower edge of
the fifth and the upper edge of the fourth scale row. All scales on
body have the borders darkened somewhat, those of the median
dorsal row visibly darker than others, making a dim median stripe ;
scales of the nine median rows discernibly lighter than the lateral
scales; edges of the ventrals with pigment, intensified more or less
into an elongate spot on outer edge. Top of head with an elaborate
scroll-like pattern; a pair of fine light spots on parietals and two
lighter-centered dark spots on each prefrontal; the dark band in
front of eyes is continuous across the end of the snout; upper edges
of supralabials with a series of black spots on the white supralabial
band; entire ventral surface immaculate.

Remarks: The nuchal collar, the head pattern, the presence of
two preoculars and the greatly reduced ventral series seem to sep-
arate this form from other known species of Rhadinaea.

Rhadinaea lachrymans has a nuchal collar but lacks the line from
the eye connecting with the lateral line, but it has fewer labials (7),
one instead of two preoculars, a higher series of ventrals (159-164)
(caudals 84-95) and a completely different body pattern.

While it differs strikingly from the head and body pattern of
Rhadinaea decorata, the reduced ventral count (128) falls into the
known range of that form (113-130; 86-123 caudals).

The species apparently belongs in the group to which R. crassa,
R. forbesi, R. gaigeae, R. quinquelineata and R. montana belong,
and its closest relationship is probably with crassa. This species
however lacks a nuchal collar and has a higher ventral count. The
general color pattern is similar but the details of the pattern differ.

Thalerophis mexicanus mexicanus Dumeril, Bibron, and Dumeril

Leptophis mexicanus Dumeril, Bibron and Dumeril, Erpetologie Generale, vol. 7, pt. 1,
1854, pp. 536-537. (Type locality, Mexico.)

Leptophis mexicanus mexicanus Oliver, Occ. Papers Mus. Zool. Univ. Michigan, no. 562,
1942, p. 10.

A series of seven specimens were obtained in the Xilitla region.

The scale formula is 15-15-11; preoculars, 1; postoculars, 2;
loreal, 1; anal, 2; 5 labials touching first chinshields. These char-
acters are invariable.

Taylor: Herpetology of San Luis Potosi, Mexico 199
Data on Thalerophis mexicanus mexicanus Dumeril, Bibron and Dumeril

No.

Locality

Sex

Ventrals

Candals

Supra-
labials

Infra-
labials

Total
length
{mm.)

Tail
length
{mm..)

251

Tamuin

9

173

164

8-8

10-10

337

123

252

Xilitla

9

175

163

8-8

10-10

984

373

253

Xilitia

$

163

167

8-8

10-10

1231

496

254

Xilitla

$

164

154

8-8

11-11

1122

255

Xilitla

9

176

164

9-8

10-10

1277

490

256

Xilitla

$

168

160

8-8

11-11

. . .

257

Xilitla

yg.

163

164

8-8

10-10

227

141

Dryadophis melanolomus veraecrucis (Stuart)

Eudryas boddaerti mexicanus Stuart, Occ. Papers Mus. Zool. Univ. Michigan, no. 254
1933, pp. 8-9. (Tj'pe locality, Zacualpan, Veracruz, Me.xico.)

Dryadophis melanolomus veraecrucis Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, no.
49, 1941, pp. 91-93, pi. 4, fig. 6, map 4 (substitute name for mexicanus).

Sixteen specimens, all from the Xilitla region, are in the collection.
The young specimens have a white line from the tip of the snout
along the lip that continues as a narrow line on the neck for a
distance equal to 16 transverse scale rows. This line is interrupted
beneath the eye by a black inverted "V." A series of white marks
are present on the lower lip, and also on many of ike chin scales,
continuing on to the first part of the ventral series where they are
larger and arranged in two rows.

Except for the labial line, some trace of this juvenile pattern can
be discerned in certain adult specimens.

The scale formula is invariable 17-17-15 in the series. There is
no variation in the following characters: preocular, 1; postoculars,
2; loreal, 1; anal, 2. The temporal scales, however, are variable.

The formula y + 2 is most frequent, but 2 + 2 occurs nearly as
frequently and 2 -f- 2 + 2 is found in four cases. Five scales are in-
variably in contact with the chinshields.

Data on Dryadophis melanolomus veraecrucis (Stuart)

Supra- Infra- Total Tail

No. Sex Ventrals Caudals labials labials length length

232 $ 178 ... 9-9 10-10

233 $ 174 94-f- 9-9 11-10 1234 396-f
247 $ 169 105 9-9 11-11 1061 324

234 yg. 180 71-|- 9-9 11-10 435 120

235 $ 176 111 9-9 11-10 1038 318

236 $ 177 103 9-9 11-11 939 284

237 $ 171 104 9-9 10-10 1100 333

238 $ 178 105 9-9 11-11 1131 330

239 $ 177 96-H 9-9 10-11 1066 314-}-

240 yg. 174 104 9-9 10-10 611 180

241 $ 173 110 9-9 11-10 1086 315

242 $ 177 9Z+ 9-9 11-10 1095 314 +

243 $ 172 98 9-9 10-10 1173 342

244 $ 176 105 9-10 11-11 1209 368

245 $ 181 ... 9-9 10-10

246 5 179 102 9-9 11-11 1095 325

200 The University Science Bulletin

Drymarchon corais erebennus (Cope)

Spilotes erebennus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860, p. 342.
(Type locality, Eagle Pass, Texas.)

Drymarchon corais erebennus Smith, Journ. Washington Acad. Sci., vol. 31, 1941, pp.
478-479, map, fig. 2.

Three specimens (nos. 248-250) were collected in the Xilitla re-
gion. The following characters are invariable in the specimens:
preocular, 1; postoculars, 2, temporals, 2 + 2; supralabials, 8;
infralabials, 8.

Data on Drymarchon corais erebennus {Cope)

No.

iSex

Ventrals

Caudals

Scale
formula

Total
length

Tail
length

248

2

182

65

19-17-14

1510

270

250

$

187

65

18-17-14

1544

298

249

5

191

65

17-17-14

1804

281

No.

Sex

Ventrals

Caudals

Supra-
labials

Infra-
labials

260

$

202

124

8-8

9-8

261

S

205

121

8-8

9-8

Spilotes pidlatus mexicanus (Laurenti)

Cerastes mexicanus Laurenti, Specimen medicum exhibens synopsin reptilium, 1768, p. 83.
(Type locality, Mexico.)

Spilotes pullatus mexicanus Amaral, Mem. Inst. Butantan, vol. 4, 1929, pp. 282-284,
fig. 2.

Two large Specimens from the Xilitla region give the following
scale data:

Data on Spilotes pullatus mexicanus (Laurenti)

Total Tail

length length

2080 565

1990 530

The preoculars and postoculars are 1 and 2, respectively, and the
scale formula is 16-18-12 in both. The coloration is normal for the
subspecies. There is a tendency for the posterior labials to fuse
with the temporals.

Pituophis deppei jani (Dumeril)

Elaphis deppei Dumeril, Mem. Acad. Inst. France, vol. 23, 1853, p. 453. (Type locality
Mexico.)

Pituophis deppei deppei Stull, Dec. Pap. Mus. Zool. Univ. Michigan, No. 250, 1932,
pp. 1-2.

A single specimen (no. 258) taken in the region near Ciudad
Maiz by Mr. Charles R. Shaw, is the first record of this subspecies
for the state. While the specimen is anomalous in certain respects,
I believe it is correctly referred to this form.

In coloration and markings it agrees reasonably well with speci-
mens from Tamaulipas listed by Stull, loc. cit. The top and sides
of the head are nearly uniform faun, with the labial region largely
cream, but with the pigment intensified on part of the supralabial
and infralabial sutures. The light spots are for the most part "en-

Taylor: Herpetology of San Luis Potosi, Mexico 201

closed" by the dark blotches, especially in the middle and posterior
parts of the body.

The prefrontal scales, however, are four instead of two, with a
small azygos scale just behind the internasal suture. There are
eight supralabials; 11-12 infralabials. The scale formula is 29-
27-21. Ventrals; 235; caudals 62; anal single; 31 body blotches,
black anteriorly, lavender posteriorly; 9 blotches on tail, darker
than those preceding.

Drymobius margaritiferus margaritiferus (Schlegel)

Herpetodryas margaritijeras Schlegel, Essai sur la Physionomie des Serpens, vol. 2, 1837,
p. 184. (Type locality unknown [stated New Orleans, in error].)

Drymobius margaritiferus margaritiferus Smith, Proc. U. S. Nat. Mus., vol. 92, 1942,
p. 383.

The scale formula of all of these specimens is 17-17-15; five
labials touch the first chinshields. The temporal formula is 2 + 2
or 2 -f 1 -|- 2. One specimen has 3 postoculars instead of the nor-
mal number of 2 on one side. This extra scale apparently is seg-
mented from the sixth labial.

Data on Drymobius margaritiferus margaritiferus {Schlegel)

No.

Locality

Sex

Ventral s

Caudals

Supra-
labials

Infra-
labials

Total
length

Tail
length

226

5.4 mi.
W. Ebano

yg-

154

120

9-8

11-11

317

105

227

Xilitla

$

148

115

9-9

10-10

729

263

228

Xilitla

$

154

9-9

11-11

. . .

. . .

229

Xilitla

$

153

122

9-8

9- 8

900

326

280

Xilitla

$

153

113 +

9-9

10-10

726

239*

231

Xilitla

9

150

109

9-9

11-11

1157

390

«

Extreme tip missing.

Lampropeltis triangulum arcifera (Werner)

Coronella micropholis arcifera Werner, Zool. Anz., vol. 26, 1903, p. 250. (Type locality,
Mexico.)

Lampropeltis triangulum arcifera Smith, Proc. Rochester Acad. Sci., vol. 8, Sept. 10,
1942, pp. 175-207.

A series of 5 specimens obtained from the Xilitla region are re-
ferred to Lampropeltis triangulum arcifera (no. 278) or are pre-
sumed intergrades between this species and L. t. polyzona (nos.
279-282).

No. 278. This specimen may be regarded as Lajnpropeltis tri-
angulum arcifera. In this the black bands are as wide or wider
than the red bands save in the first. Fifteen of the red bands are
interrupted by black on the medial line. All red bands save first
on tail are interrupted or suppressed completely. The nuchal red
band is much reduced, its width being equivalent to five scale
lengths.

202 The University Science Bulletin

No. 279 has a number of the dark bands reaching a width equal
to that of the red bands and toward the posterior part of the body
one or two of the black bands are separated by less than the length
of one red scale. The tail has six white bands, and three of the
black tail bands are split with red.

No. 280 is similar to this, but more of the red bands are wider
than the black ones; three of the black bands are split with red.

No. 281 is similar to the preceding but there are eight white bands
on the tail, and only one is partially split with red. The black and
red bands are more nearly equal in width. Anteriorly, however,
the red bands are wider.

In no. 282, many of the black bands meet (thus dividing the
red band) or they are narrowly separated. There are five white
bands on the tail, and the red is suppressed except on two of the
black bands.

The exact localities for these snakes is not available, the labels
bearing the notation Xilitla region. It is not known whether the
t. polyzona and t. arcifera were taken in the same exact locality
or whether they were separated by some considerable difference in
altitude.

Data on Lampropeltis t. arcifera and intergrades

No.

Sex

Scale
formula

Ventrals

Caudals

White
body
ba7ids

Tail
bands

Temporals

Total

length

Tail

length

281

$

21-21-19

213

58

22

8

2-1-3

976

125

282

$

21-21-17

213

51

22

5

2 -t- 3

1010

152

279

S

22-21-17

215

49-f-

23

5

2 -f 2

1176

1554-

280

9

22-21-19

208

60

18

5

2 + 2 (2 -fl)

682

115

278

?

22-21-19

208

55 +

25

8

2 -h 3 (2 + 3)

540

85 +

Lampropeltis triangulum polyzona Cope

Lampropeltis polyzona Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860, p. 258.
(Type locality, Cuatupe, near Jalapa, Veracruz.)

Lampropeltis triangulum polyzona Dunn, Gee. Pap. Mus. Zool. Univ. Michigan, no. 353,
1937, p. 1.

Four specimens from the Xilitla region are referred to this sub-
species (nos. 274-277). They are in general agreement as to the
color pattern. This consists of black-white-black bands, separated
by red bands. In each case the red bands are wider, usually much
wider, than the bordering bands. The black bands are somewhat
wider medially than on the sides or ventrally. All of the specimens
have the scale formula 21-21-17.

The anterior red bands are eight to ten scale lengths wide; the
light bands usually one and one-half to one scale length wide. All
the scales of the red bands are black-tipped and the same is true of
the scales of the white bands. On the venter the white bands oc-

Taylor: Heepetology of San Luis Potosi, Mexico 203

cupy one or two ventrals; when only one ventral is covered, some
parts of the adjoining ventrals may be white.

The specimens have from 18 to 20 white bands on the body, and
four to six white bands on the tail (possibly all with six, since the
tail tips are missing in three). Three of the black tail bands are
"split with red" of varying width. The ventrals may show darker
edges or there may be flecks of black on both red and white scales.

One might expect the range of Lampropeltis trianguluni annulata
to extend south from Tamaulipas and Nuevo Leon into the northern
and eastern part of San Luis Potosi, but as yet none has been found
referable to this subspecies.

Data on Lampropeltis t. ■polyzona (Cope)

No.

Sex

Ventrals

Caudals

White

body

bands

White

tail

bands

Temporals

Total
length

Tail
length

274

$

220

52 +

18

5

2 + 2 + 3

845

112 +

275

$

213

47 +

20

^ 4

2 + 1 + 3

746

132 +

276

$

212

50 +

20

4

2 + 3

975

138 +

277

$

214

63

19

6

2 + 3

927

158

Elaphe fiavirufa (Cope)

Coluber flavirufiis Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18, 1866 (1867), p.
319. (Type locality Yucatan.)

Elaphe fiavirufa fiavirufa Smith, Copeia, 1941, no. 3, p. 132, fig. 2.

Two specimens, no. 271 $ and no. 272 5 , obtained in the Xilitla
region, are the first state records.

Scale data for the two, respectively, are as follows: scale formula,
26-29-21, 27-31-23; ventrals, 254, 253; caudals, 108, 105; suprala-
bials, 9-9, 10-9; infralabials, 12-12, 13-14; preoculars, 1-1, 1-1,
postoculars, 2-2, 2-2; temporals (somewhat irregular), 3 + 4, 2
4- 3; spots on body, 36, 37; spots on tail, 18, 17; total length (in
mm.) , 632, 1242 ; tail, 134, 265.

Elaphe laeta laeta (Baird and Girard)

Scotophis laetus Baird and Girard, Catalogue of North American Reptiles, 1853, pp. 77-78.
(Type locality, Red River, Arkansas.)

Elaphe laeta laeta Woodbury and Woodbury, Proc. Biol. Soc. Washington, vol. 55, 1942,
pp. 139-140.

This specimen (no. 273), the first recorded for the state, was
taken at Ebano in the extreme northeastern part. It is a female
having 230 ventrals and 70 caudals. The scale formula is 25-27-19 ;
the supralabials and infralabials are 8 and 10 respectively. There
are 41 body blotches and 17 blotches on the tail, the most posterior
ones being too indistinct to be counted. One preocular and 2 post-
oculars are present. The total length is 437 mm.; the tail, 71 mm.

204 The University Science Bulletin

Pliocercus laticollaris Smith

Pliocercus elapoides laticollaris Smith, Proc. Biol. Soc. Washington, vol. 54, 1941, pp.
122-123, and vol. 55, 1942, p. 160.

The finding of a specimen of this rare snake at Xilitla by Charles
R. Shaw traces its distribution to the northwest nearly three hun-
dred miles, and adds an interesting species to the fauna of the state.

There is a white band, bearing a fine peppering of pigment, that
crosses the back of the head involving all of the parietals save their
anterior edges, and including the anterior part of the first scale row
following the parietals. The band is continuous with the white color
which covers the chin, the lower lip and most of the upper lip. The
rest of the head is coal-black.

The body pattern consists of yellow or cream-edged black rings.
The black covers from 3 to 3^2 scale rows, the cream usually only
one-half scale row. The intervening spaces are red in color, forming
bands around the body that cover six transverse scale rows. Each
red scale bears a black dot.

The black bands are incomplete in the region anterior to the anus,
and this space on the venter is creamy white involving parts of four
ventrals. Altogether, there are 16 black body bands (including the
nuchal) and 15 red bands. On the tail the black bands have wider
cream borders and are continuous around the tail ; but the last five
are reddish dorsally, with rather heavy, dark flecks, and cream on
the ventral side.

Scale data: ventrals, 136; caudals, 85; preoculars, 2-1, the lower
on the right side small, wedged in between third and fourth labials;
supralabials, 8-8; infralabials, 8-8; total length, 541 mm.; tail, 193
mm.

This specimen differs from the southern (Tabasco) specimens in
having eight instead of ten labials both above and below. The wide
distribution of laticollaris with at least two other forms intervening
in the territory, elapoides elapoides and elapoides celatus, strongly
suggests that the form should be regarded as a species rather than as
a subspecies.

Thamnophis scalaris scaliger (Jan)

Tropidonotus scaliyer Jan, Elenco sistematico degli Ofidi, 1863, p. 70. (Type locality-
unknown.)

Thamnophis scalaris scaliger Smith, Zoologica, vol. 27, 1942, pp. 103-104; Smith and
Taylor, U. S. Nat. Mus. Bull., no. 187, Oct. 5, 1945, p. 167 (doubtful reference in San Luis
Potosi).

A series of specimens, taken in the general region of Xilitla and
Cerro Cone jo, are referred to this subspecies.

In four specimens the scale formula is 19-19-17, one only having

No.

Sex

291

9

292

$

293

9

294

9

Taylor: Herpetology of San Luis PotosI, Mexico 205

16 scale rows anterior to the vent. The upper labials are 8-8, save
in one where the number seven occurs on one side. The lower labials
are 10-10, with two exceptions; in these nine are present on one side.
The ventral averages are somewhat higher than those given by
Smith {loc. cit., p. 103). The ventrals are 151-162; the caudals, 71-
72, in males. The ventrals are 151-157; the caudals, 65-66, in fe-
males. There is an increase of one in the number of upper labials.

Data on Thamnophis scalaris scaliger (Jan)

Locality Ventrals Caudals Preoculars Postoculars Temporals Length Length

Cerro Conejo 157 66 1-1 3-3 1 + 2

CerroConejo 162 71 1-1 3-3 1 -f 2 676 102 +

Xilitla 154 65 1-1 3-3 1 + 2 612 129

Xilitia 151 41+ 1-1 3-3 1 + 2 676 102 +

Thamnophis phenax phenax (Cope)

Eutaenia phenax Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 20, 1868, p. 134. (Type
locality, Cordoba, Veracruz, Mexico.)

Thamnophis phenax phenax Smith, Zoologica, vol. 27, 1942, pp. 99-100.

Four specimens of this rare snake have been taken in the Xilitla
region. One (no. 284) bears the notation "Miramar 4,500 ft."

The scale formula is 19-19-17 in three specimens; one specimen
has the rows reduced to 15 in front of vent; supralabials, 8-8; infra-
labials, 10-10, or 9-10; preoculars, 1; postoculars, 3. The temporals
are 1 -f- 2 (or 1 -)- 3 in a single instance) ; loreal, 1 ; anal single. All
show the pattern of large quadrangular blotches.

No. 285 contained the remains of a small terrestrial salamander,
which is described elsewhere in this paper.

No. 283 has a median white line dividing the anterior spots, which
tend to alternate.

Data on Thamnophis phenax phenax (Cope)

No.

Sex

Locality

Ventrals

Caudals

Body
blotches

Total
length

Tail
length

283

S

Xilitla

151

72

41

548

131

284

9

Xilitla, Miramar

4,500 ft.

159

62

37

548

105.5

285

yg-

Xilitla

159

75

48

157

48

286

yg.

Xilitla

160

80

41

163

51

287

s

161

78

41

Thamnophis marciana (Baird and Girard)

Eutaenia marciana Baird and Girard, Catalogue of North American Reptiles, 1853, pp.
36-37. (Type locality, restricted to Red River, Arkansas. Originally included New Braun-
fels, San Pedro, and Indianola, Texas.)

Thamnophis marciana Ruthven, U. S. Nat. Mus. Bull., 61, 1908, pp. 849-852, pi. 93.

Smith and Taylor * have regarded a record of this species for San
Luis Potosi ** as doubtful. The finding of three specimens in the

* U. S. Nat. Mus. Bull., no. 187, Oct. 5, 1945, p. 164.
*» Carman, Bull. Essex Inst., vol. 19, 1887, pp. 7-8.

206 The University Science Bulletin

northeastern part of the state by Mr. Charles R. Shaw and Marcella
Newman establishes the species beyond question as a resident of the
lowlands in the eastern part of the state. Two were taken Novem-
ber 6, 1946, and one on March 19, 1947. The following data were
recorded:

The following scale characters are invariable: scale formula,
21-21-17; supralabials, 8-8; infralabials, 10-10; loreal, 1; anal, 1;
5 scales touching first chinshield. The temporals are 1 -j- 2, 1 + 3,
or 2 + 2.

The lateral light stripe is restricted to the third scale row. A verte-
bral light (orange) stripe one whole, and two half scale rows wide,
is evident the length of the body.

Data on Thamnophis marciana {Baird and Girard)

No.

Sex

Locality

Ventrals

Caudals Preoculars Postoculars

Total
length

Tail
length

288

$

Ebano

149

62 1-1 4-4

335

71

289

9

Ebano

146

58 1-1 4-4

408

89

290

$

Sabanito

151

69 1-1 3-4

460

110

Storeria dekayi texana Trapido

Storeria dekayi texana Trapido, Amer. Mid. Nat., vol. 31, no. 1, pp. 63-69, figs. 51-60.
(Type locality, San Rafael, Jicaltepec, Veracruz, Mexico, alt. circa 100 ft.)

Two specimens from the Xilitla region are referred to this re-
cently described form. The species has been reported from the state
on the basis of paratypic * specimens which I collected, 5 miles
south of Valles.

Data on Storena dekayi texana Trapido

No.

Sex

Locality

Ventrals

Caudals

Supra-
labials

Infra-
labials

Total
length

Tail
length

268

5

Xilitla

141

47

7-7

7-7

350

64

269

5

Xilitla'

141

364-

7-7

7-6

322

60 -h

The scale formula is 17-17-17; postoculars, 2-2; preoculars, 1-1;
temporals, 1 + 2, and 1 + 2, 1 + 3.

Seven of the subcaudals of no. 269 are entire, while the remainder
are normally divided; the fifth and fourth labials are fused together.
The adult markings are much obscured.

No. 269 contains 8 nearly full-time embryos, which average about
84 mm. in length. Paired spots on the dorsum are evident.

No. 268 contains several eggs, but in none did I discern embryos.
This specimen was captured May 13; the other with the embryos
was taken June 29.

It is significant that there are two subspecies of Storeria dekayi
reported from San Luis Potosi. The two specimens of Storeria

* These are not specifically designated as paratypes but were dealt with in the type de-
scription.

Taylor: Herpetology of San Luis Potosi, Mexico 207

dekayi texana mentioned in the type description "5 miles south of
Valles, District Ciiidad de Valles EHT-HMS 4662, 2664" were
taken in the same pile of driftwood in which was found Storeria
dekayi tem'poralineata mentioned in the type description of that
form as "five miles south of Valles, EHT-HMS 4663." If these are
distinct, one might question the wisdom of regarding them as sub-
species. Sufficient material is not at hand to determine this point.

Tantilla rubra Cope

Tantilla rubra Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876. (Type
locality, "Tapanatepec" = Tapana, Oaxaca) ; Smith, Zoologica, vol. 27, 1942, p. 40.

One specimen from the Xilitla region (no. 305 J ) has the head
missing save for certain fragments of skin and one lower jaw.

The scale formula is 15-15-15. The mental is separated from the
chinshields. There are 7-7 infralabials, the first four in contact with
the first pair of chinshields. There are 157 ventrals, and 64 caudals.
The total length (length of head estimated) 345 mm.; length of tail,
85 mm.

The general color is a pinkish brown, the edges of each scale
somewhat darker than the center. Ventrals and caudals are pink.
The outer half of outer scale row is cream, with a slight wash of
pink, scarcely differentiated from the ventral coloration. The upper
half of the outer scale row is distinctly darker than adjacent rows.
On the tail the lower half of the outer scale row has a small trian-
gular blackish spot on each scale.

A broad, black, nuchal collar is present, three to four scales in
width. Preceding this is a cream collar two and one half scale rows
wide. A black occipital collar crosses the parietals and reaches to
the supralabial edges. A few black spots on infralabials. Other
details of the head coloration are wanting.

Tantilla shawi sp. nov.

Type: Louisiana State University Museum no. 306. Collected at
Xilitla (Miramar) , Aug. 28, 1947, by Charles Shaw.

Diagnosis: A tantilla characterized by seven supralabials, six
infralabials, the chinshields separated from the mental; ventrals,
169; subcaudals, 48; anal divided.

Tip of snout black, bordered by a cream band that is followed by
a broad black area reaching to the back part of parietals; cream
spot behind eye extending from frontal to labials ; an occipital cream
band one scale wide, followed by a broad nuchal black band four

208 The University Science Bulletin

scales wide. Anterior fourth of body banded with narrow cream
and wider black bands.

Description of type: Rostral broader than high, visible above as
a very narrow triangle, the summit curved rather than angular;
internasals a little less than half area of prefrontals, which are
very broad, in contact laterally with second labial; frontal hexa-
gonal, obtusely angled in front, acute behind; distinctly longer than
wide (1^/4 times), its length greater than its distance from tip of
snout, its width not quite twice that of supraoculars; parietals mi-
nutely longer than their distance from tip of snout; nasal divided,
the anterior part about a half larger than the posterior, and sep-
arated from preocular by prefrontal ; nostril between two nasals, the
posterior widely separated from preocular ; one preocular, two post-
oculars; a large anterior temporal followed by a posterior temporal
that is somewhat smaller; supralabials, 7-7, in following order of
size: 1, 3, 2, 5, 4, 6, 7, the third and fourth border the orbit; 6-6
infralabials, the first pair separating mental and chinshields; four
labials touching the first chinshields, which are nearly three times
area of second pair; latter pair in contact; diameter of eye twice
in snout length, and one and one-fourth times in eye to nostril dis-
tance. Scale formula, 15-15-15; the scales smooth without trace of
apical pits; ventrals, 169; caudals, 48; anal divided; total length,
369; tail, 73.5.

Color: Tip of snout blackish, bordered by a cream band that
crosses snout, reaching mouth; beginning on the anterior part of the
prefrontals and extending to near the posterior tip of parietals, a
broad black band reaching on side of head to mouth ; a cream spot,
covering part of the fourth and fifth supralabials, extends up some
distance behind the eye, reaching the frontal; an occipital cream
band about one scale wide connects with cream color of chin and
throat; this followed by a broad nuchal dark band about three and
one half scales wide. The infralabials and mental with variable-
sized areas of black, bordered partially by cream; the spot on the
fourtli infralabial reaches to chinshield; that on fifth small, leaving
a large cream area.

General color of body black or bluish black, the edges of the
scales faintly outlined in cream or white; scales of outer row with
widest light edges; a series of small black dots on outer edge of ven-
trals (except first 18 or 20) and caudals; venter and subcaudal
region light pink; chin and throat cream; anterior fourth of body
traversed by narrow transverse cream lines, usually less than one
scale wide, and which may be broken mesially and tending to alter-

Taylor: Herpetology of San Luis Potosi, Mexico 209

uate with that from opposite side. There is a suggestion of a dis-
continuous median line on this part of body.

Relationship: The species differs from all other Mexican species
in the character of the annulation of the anterior fourth of the body,
and in the details of the head markings.

The Central American Tantilla annulata has this tendency to
annulation carried still farther back, some of the semiannulations
being present on the tail. That form, however, is one of the largest
in the genus — a Costa Rican specimen examined having a total
length of 502 mm., the tail being 108 mm. The character of the
markings, yellow bars with black borders, is such that there is no
possibility of confusing the species with Tantilla shawl.

The species is named for Mr. Charles Shaw, its discoverer.

Coniophanes imperialis imperialis (Kennicott)

Taeniophis imperialis Girard, The U. S. Naval Astronomical Expedition to the Southprn
Hemisphere during the years 1849-'50-'51-'52, Reptiles, Senate Doc, no. 121, 1855, p. 216.
(Type locality, Matamoros, Tamaulipas, Mexico.)

Coniophanes imperialis imperialis Cope, Ann. Rept. U. S. Nat. Mus., 1898 (1900), p.
1090; Bailey, Papers Michigan Acad. Sci. Arts Lett., vol. 24, pi. 2, 1938 (1939), pp. 34-36,
pi. 1, fig. 1, map, fig. 5.

Two specimens (nos. 296, 297) were taken on the lowlands at
Ebano in the extreme northeastern part of the state. These agree
in the following characters: scale formula, 19-17-15; supralabials,
8-8; infralabials, 9-9; preoculars, 1-1; postoculars, 2-2; temporals,
1 + 2; loreal, 1; anal, 2.

The ventral scale count and measurement in mm. for 296 is 135;
caudals 71; total length 398; tail 111; for 297, ventrals 132; caudals
74; total length 140, tail 44. Both are male.

From Bailey's diagnosis of this form, loc. cit., these differ in hav-
ing 17 scale rows about the middle of the body instead of 19. They
agree, however, rather completely in the characteristic color pattern
shown in Bailey's illustrations, loc. cit.

In 297, a recently hatched specimen, the markings are very
sharply defined. The light line through the eye is white, edged with
black, and extends along the canthal region and across the snout.
The anterior end of the dorsolateral stripes are white, but soon
develop posteriorly a scattering of brownish pigment. The lower
edge of the white line is bordered by a black line, which is the upper
edge of the broad lateral dark stripe that covers 4% scale rows.
The supralabials are bordered above by a black line, below which is
a white line. Below this line the lower part of the labials are flecked
v/ith black pigment. The chin and infralabials are also flecked with

14^3659

210 The University Science Bulletin

pigment, and there is a distinct spot on the fifth infralabial. There
is a very indefinite pigmented area on the outer edges of the ven-
trals.

There is no apparent approach to Coniophanes imperialis clavatus
or proterops, forms that occur to the south.

These specimens are the first records of the subspecies for the
state.

Coniphanes fissidens proterops (Cope)

Cloniophanes] proterops Cope, Proc. Acad. Nat. Scl. Philadelphia, vol. 12, 1860, p. 249.
(Type locality, vicinity of Jalapa, Veracruz.)

Coniophanes fissidens proterops Smith, Proc. U. S. Nat. Mus., vol. 91, 1941, pp. 107-109,
map, fig. 33.

A single specimen of this species (no. 295) is in the collection from
the Xilitia region. It is the first record of the form for San Luis
Potosi.

The specimen shows the following characters: scale formula, 19-
19-17; ventrals, 129; caudals, 30-1- (tail broken); supralabials, 7,
infralabials, 10 ; preocular, 1 ; postoculars, 2 ; temporals, 1 -|- 2 ; 5
labials touch first chinshields; length of body 300; tail partially
missing.

The median dorsal "black" line consists only of the dark edges of
the median dorsal scale row, and taken together the spots have the
appearance of a chain.

Trimorphodon tau Cope

Trimorphodon tau Cope, Proc. Amer. Philos. Soc, vol. 11, 18G9, pp. 151-152. (Type
locality Quiotepec, Oaxaca) ; Taylor, Univ. Kansas Sci. Bull., vol. 26, 1939 (1940), pp.
474-477, fig. 8, pi. 51.

A single specimen of this species (no. 298), captured two and one-
half miles south of Pendencia, San Luis Potosi, December 2, 1946,
traces the known range from the southern edge of the plateau, a
distance of some three hundred miles. Heretofore the species has
been known from possibly half a dozen specimens, with localities in
Michoacan, Guerrero, and Oaxaca.

This specimen has been compared with Oaxaca specimens and
while a few differences exist it does not appear to merit nomencla-
tural designation. A larger series from this region might point to a
separation.

The specimen is a female having a scale formula 23-21-15
(Oaxaca specimens have 23-20-16 and 22-21-14); ventrals, 218;
caudals, 59; anal, 2; supralabials, 8-8; infralabials, 11-11; pre-
oculars, 3-3; postoculars, 3-3; loreals, 3-2; temporals, 2 + 3-1-4, 2
+ 2 -|- 4, somewhat irregularly placed; four infralabials touch chin-

Taylor: Herpetology of San Luis Potosi, Mexico 211

shields; anterior chinshield double the size of posterior; fourth and
fifth labials enter orbit; spots on body, 27; 12 or 13 on tail.

The markings of the head and general markings of the body
approximate rather closely the pattern depicted by Taylor, fig. 8,
loc. cit., and pi. 51. However, the ground color of the specimen is a
little darker. The edges of the ventrals have indistinct darker spots
which are the ends of the dorsal blotches and small spots that alter-
nate with them. There may be some peppering of pigment on the
ventrals. scarcely visible to the naked eye. There are a few indefi-
nite dark marks on the chin.

The following shows the similarity in scale counts between this
specimen and two in the EHT-HMS collection (nos. 5506, 5507
from San Felipe, Oaxaca, near the city of Oaxaca). LSU no. 298 5
ventrals, 218; subcaudals, 59; total, 277. No. 5506, ^, ventrals,
206; subcaudals, 70; total, 276. No. 5507, $ , ventrals, 220; sub-
caudals, 57; total 277.

Leptodeira maculata (Hallowell)

Megalops maculatus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, 1860 (1861),

p. 468. (Type locality, "Tahiti," by error.)

Leptodeira maculata Taylor, Univ. Kansas Sci. Bull., vol. 25, 1938 (1939), pp. 337-342,

figs. 6-7, pi. 31, fig. 1; pi. 32; pi. 33, figs. 1-3.

One specimen (no. 299) was taken near Sabanito, El Salto Junc-
tion, Naranja Region, San Luis Potosi. The* ventrals are 169, the
subcaudals 59. Other scale data are: preoculars, 2, postoculars,
3-2; upper labials, 8; lower labials, 10; 26 body spots, 8 caudal spots
reaching laterally to the second scale row; scale formula, 23-25-17.
The characteristic head marking consists of a white border on the
frontal scale, connecting with a line foraied by the white edges of
the parietals that follows their common suture.

The specimen was taken in "palm jungle by a marsh."

Leptodeira annulata septentnonalis (Kennicott)

Dipsas septentrionalis Kennicott, in Baird, Report of the U. S. Mexican Boundary Survey,
vol. 2, 1859, Reptiles, p. 16, pi. 8, fig. 1. (Type locality, Matamoros, Tamaulipas, Mexico,
and Brownsville, Te.xas, U. S. A.)

Leptodeira annulata septentrionalis Smith, Proc. Biol. Soc. Washington, vol. 54, 1941,
p. 117.

Six specimens were taken, five from the Xilitla region and one
from the extreme northeastern part of the state at Ebano.

The series shows complete unifomiity in the following characters:
scale formula, 21-23-17; supralabials, 8; infralabials, 10; preoculars,
3; postoculars, 2; scales touching first chinshields, 5.

Smith, loc. cit., has recently described a new form, Leptodeira

212 The University Science Bulletin

annulata taylori, from Orizaba, Veracruz, characterized by 196-206
ventrals; 80 to 101 caudals; usually three preoculars, and spots not
extending below the third scale row; from 36 to 47 dorsal spots on
body.

In the series at hand the spots reach either the first scale row (3
specimens), the second row (one specimen), or the third row (one
specimen) . The northern septentrionalis ordinarily reduce the scale
rows to 15 in front of the anus and the spots are usually less than
25 — rarely above thirty.

Despite certain differences it seems that the affinities of these
specimens are closer to L. a. septentrionalis than to L. a. taylori, and
not impossibly they represent a population similar to that at Tux-
pan which Smith, loc. cit., regarded as representing intergrades.

The stomach of no. 300 contained the remains of an unidentified
Sceloporus. No. 302 had 9 eggs in the oviducts.

Data on Leptodeira annulata septentrionalis (Kennicott)

No.

Sex

Locality

Ventrals

Caudals

Body
spots

Tail
spots

Total
length

Tail
length

300

$

Eliano

195

83

30

15

386

87

301

yg.

Xilitia

193

94

28

18

420

106

302

9

Xilitia

193

77

29

12

825

158

303

9

Xilitia

205

75

29

14

820

156

304

S

Xilitia

206

66

32

14

818

146

Micruriis fitzingeri microgalbineiis Brown and Smith

Micrurus fitzingeri microgalbineus Brown and Smith, Proc. Biol. Soc. Washingi;on, vol.
55, 1942, pp. 63-67. (Type locality, 7 km. south of Antiguo Morelos, Tamaulipas, Mexico.)

The collection contains three specimens of this form which has
been known previously from only two specimens, one from the type
locality and one from Tamuzunchale, San Luis Potosi.

The following data add considerably to our knowledge of the
variation in subspecific characters:

Scale formula: 15-15-15; supralabials and infralabials, 8. Pre-
oculars, 1; postoculars, 2; four scales touch the chinshields. Tem-
porals 1 + 1 + 2 in two specimens, 1 + 1 + 1 in the other.

The color pattern is formed by a series of alternating black and
red bands, separated by very narrow white bands. Usually the
white part consists of separate white spots covering about one half
of a scale and not or barely contiguous with the next white spot. As
a result of this the red bands appear to have a zigzag edge, while the
black bands present an edge that is nearly straight.

The black bands usually cover from three to four scale rows;
occasional bands cover only 2^/^ scale rows. The intervening red
bands vary from 6 to 7 scale rows in length. More than half of the

Taylor: Herpetology of San Luis Potosi, Mexico 213

scales of all the rows except the outer have a black spot. The black
spots vary much in size, being largest about the middle of the body.
A few black spots usually smaller than a dorsal scale are scattered
sparsely and irregularly on the ventrals.

The white band crossing the parietal region is half or a little less
than half the length of the parietal. It passes somewhat forward
involving the entire sixth labial and parts of the fifth and seventh
labials. The rest of the upper labials are intensely black. Three
white areas present on the chin, the two outer covering parts of the
fourth and fifth lower labials, the median, the anterior part of the
second pair of chinshields. In one specimen the spots are practically
contiguous, thus forming a band entirely around the head.

The female specimen no. 309 has five black and four white bands
on the tail, the black being 6 to 6^/2 scale lengths wide, the white 2
scale rows wide. The males have 6 or 7 black bands on the tail of
about the samg length as those in the females. The white bands in
no. 307 are partially red below and above, and some of the scales
bear black spots. No. 308 shows only a faint shade of red on one
or two of the white scales.

A specimen (no. 310) of Micrurus is referred to this form with con-
siderable reluctance. The white band across the parietals is wider
(nearly % the width of the parietals) involving all the fifth and
sixth supralabials and part of the seventh, then passing across the
chin involving 5th and 6th infralabials, and most of both pairs of
chinshields.

The black blotches on the body are a little wider and occupy four
ventrals below, more often than three. The intervening red areas
have practically every scale bearing a black fleck, and on the ven-
trals there is usually a pair of heavy black spots much larger than
those on the dorsal part of the red blotches. The neck and body has
20 black blotches, the tail four. There are four white bands on the
tail.

The scale data agree with those given for the Xilitla specimens
save that ventrals are 222, the caudals 32. The black bands have
zigzag rather than straight edges. The snake is 703 mm. in length;
the tail 70.

This specimen comes from 17 km. west of Ebano in the north-
eastern lowlands of the state. It will be noted that the character of
the head band suggests the condition that obtains in the typical sub-
species M. f. fitzingeri. That form is known, however, only from
the high plateau.

214 The University Science Bulletin

Data on Micrurus fitzingeri micro galbineus Brown and Smith

Body Tail Total Tail

No. Sex Locality Ventrals Caudals bands bands length length

307 $ Xilitla 204 44 24 7 732 102

308 $ Xilitla 208 42 27 6 735 93

309 2 Xilitla 220 38 25 5 G25 65

Bothrops nummifer (Riippell)

Atropos nummifer Riippell, Verz. Mus. Senck., Amph., 1845, p. 21. (Type locality
Mexico.)

Bothrops nummifer Jan, Elenco sistematico degli Ofidi, 1803, p. 126; Smith, Proc. U. S.
Nat. Mus., vol. 93, 1943, pp. 398-401.

There are two specimens of Bothrops, nos. 318, 319, taken at
Xilitla June 19, 1947, that are puzzling, inasmuch as they occur at
the extreme northern part of the range of Bothrops nummifer (as in-
terpreted by Smith, loc. cit., yet they agree in having certain general
characteristics of B. mexicanus which tend to separate that form
from B. nummijer.

In ventral and subcaudal counts the specimens appa'oach the con-
dition in the northern B. nummijer. In the scale rows about the
body, one approaches B. nummijer, one the southern B. mexicanus.
In the character of the rostral (in contact with the nasal, first
labial and two other scales laterally, and with a small median post-
rostral) they should be referred to mexicanus. In dorsal pattern
they are more or less intermediate, one approaching the character
of mexicanus, the other, as regards the fusing of the dorsal blotches
into a zigzag or undulating band, is quite like nummijer.

However, I feel that a larger series of specimens should be ex-
amined before it is decided whether or not this is an "intergrading"
population.

No. 318 contained a ''1/3 grown black rat."

Data

on

Bothrops nummijer {R

uppell)

No.

Sex

Scale
formula

Ventrals Caudals

Supra -
labials

Infra-
labials

Tolal
length

Tail
length

318

$

25-25-20

134 33

10-9

12-11

661

75

319

S

23-23-19

131* 33

9-9

11-10

520

62

Bothrops atrox asper (Garman)

Bothrops atrox Dumeril, Bibron and Dumeril, Erpetologie generale, vol. 7, pi. 2, 1854,
pp. 1507-1509. (Type locality, Obispo, Panama.)

Bothrops atrox asper Smith and Taylor, Bull. U. S. Nat. Mus., no. 187, 1945, pp. 180-181.

Seven specimens were taken, three at El Salto, three at Xilitla.
The subcaudals in all are in a double row. One or two scale rows
intervene between the long subocular and the labials. The number
of loreals varies between three and four. There are two preoculars,
two postoculars, and usually 10 or 11 scales between the supra-

Taylor: Herpetology of San Luis PoTOsi, Mexico 215

oculars. No. 311 has the first pair of infralabials broken, forming an
extra pair of "chinshields."

Data on Bothrops atrox asper (Garman)

No.

Locality

Sex

Scale
formula

Ventrals

Caudals

Supra-
labials

Infra-
labials

Total
length

Tail
length

311

El Salto

5

27-28-19

205

62

7-8

11-10

789

109

312

Xilitla

27-25-20

203

63

7-7

10-11

1282

176

313

Xilitia

$

25-25-20

205

70

7-7

11-11

1035

150

314

Xilitla

25-29-18

203

63

7-8

10-11

1025

127

315

EI Salto

28-29-21

209

63

7-8

10-11

890

112

316

El Salto

$

28-25-21

210

67

7-7

10-10

854

120

317

$

28-25-21

204

68

7-7

10-10

1410

195

Crotalus triseriatus triseriatus Wagler

Urosophus triseriatus Wagler, Naturliches System der Amphibien, 1830, p. 176. (Type
locality Mexico.)

Crotalus triseriatus triseriatus Klauber in Githens and George, Bull. Antiv. Inst. America,
vol. 5, 1931, p. 33 (part); Gloyd, Chicago Acad. Sci. Spec. Publ., no. 4, 1940, pp. 84-91,
map 6, pi. 6, fig. 1, 2; pi. 31, fig. 4 (reported from Alvarez, San Luis Potosi).

Ten specimens of Crotalus in the collection are referred to this
species. Of the ten specimens, eight are from the Xilitla region, at
an elevation from about 3,800 feet to 4,400 feet; two are from Cerro
Conejo at a somewhat higher elevation. For the most part, the spe-
cimens are uniform in color and markings, and agree in having a
series of from 36 to 44 quadrangular blotches on the body, and from
5 to 7 on the tail. However, one of the series (no. 321) differs
markedly in having the head and body an almost uniform, slaty
black. On the posterior sixth of the back, a few very dim lighter
narrow marks can be discerned with difficulty when the specimen is
held in the proper light.

The ventrals are less intensely dark than the other specimens and
have lighter edges. The tail has some dim orange coloration ven-
trally and laterally. The keels on the scales are seemingly less
strongly elevated than those on the remainder of the series. The
scale counts of the specimen, however, fall within the known varia-
tion of the species. No melanistic specimens have heretofore been
reported, although certain other aberrant specimens have been cited
by Gloyd, loc. cit. This specimen contained a partially digested
salamander in its stomach and very numerous tapeworms.

The ventral scale series, whether male or female, averages 154.
This is the average given by Gloyd for the San Luis Potosi speci-
mens.

One specimen (no. 325) presents a curious anomaly, in having the
lower preocular completely absent. As a consequence, the orbit and
the sensory pit are confluent. The pit itself is divided into two
compartments by a heavy membrane. This condition obtains on
both sides of the head.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. I] April 20, 1949 [No.

A Review of the Lizard Genus Barisia

BY

J. A. TlHEN

Abstract. — The generic name Barisia is revived for a group of lizards that
in recent years has been placed in the genus Gerrhonotus. The following spe-
cies and subspecies are recognized: Barisia moreleti moreleti, B. m. temporalis,
B. VI. Salvador ensis, B. m. julva, B. m. rafaeli, B. monticola, B. viridiflava, B.
gadovii gadovii, B. g. levigata, B. niodesta, B. antauges, B. rudicollis, B. levicol-
lis, B. imbricata imbricata, B. i. ciliaris, B. i. planifrons.

The various groupings within the genus are discussed. Brief descriptions
are given of all the forms with their present known distribution.

A review of the lizard genus Barisia

BY

J. A. TlHEN

The genus Barisia was proposed by Gray in 1838 to contain three
species formerly included in the genus Gerrhonotus Wiegmann;
these species were B. rudicollis, B. imhricata, and B. lichenigera; B.
adspersa was included as a synonym of the last-named species. The
same arrangement appeared in his 1845 Catalogue, but he there
changed the spelling from Barisia to Barissia; this emended spelling
has incorrectly been used by all subsequent authors.

This genus has been recognized as valid by some authorities, rele-
gated to the synonymy of Gerrhonotus by others, and considered as
a subgenus of Gerrhonotus by yet others. It was redefined by Cope
in 1877 on the basis of the arrangement of the "internasal" and
frontonasal elements. He recognized five species: antauges, bo-
courti, lichenigera, imbricata, and rudicollis. In the same work he
proposed the genus Mesaspis for two forms, moreleti and julvus.
Most authors have never considered this latter a valid genus, al-

(217)

218 The University Science Bulletin

though Cope still recognized it in 1900; recent workers have also
tended to return Barisia to the synonymy of Gerrhonotus.

I have recently * set forth my reasons for believing that Barisia
should be considered a valid and recognized genus of gerrhonotine
lizards, and have redefined the genus as shown below. A discussion
of the genus as a whole, in relation to other gerrhonotine genera, is
to be found in the previous paper; the present paper deals with the
forms and interrelationships within the genus.

I wish to express my appreciation for the many helpful sugges-
tions and criticisms offered during the course of my study and the
preparation of this paper by Dr. S. C Bishop of the University of
Rochester and Dr. Hobart M. Smith of the University of Illinois,
also by Dr. E. R. Dunn of Haverford College and Drs. L. C. Stuart
and Norman Hartweg of the University of Michigan. The following
individuals have been very generous in lending personal specimens,
or specimens under their care in the institutions with which they are
connected: Dr. Doris M. Cochran of the U. S. National Museum,
Mr. Charles M. Bogert of the American Museum of Natural His-
tory; Mrs. Helen T. Gaige and Dr. Norman Hartweg of the Univer-
sity of Michigan, Dr. Edward H. Taylor of the University of Kan-
sas, Mr. Karl P. Schmidt of the Chicago Natural History Museum,
Sr. Rafael Martin del Campo of the Institute de Biologia of Mexico,
Mr. R. T. Moore of the California Institute of Technology, Mr.
Arthur Loveridge of the Museum of Comparative Zoology, and Dr.
E. R. Dunn of the Philadelphia Academy of Natural Sciences.

The following abbreviations will be used throughout this paper in
reference to the various collections examined:

AMNH American Museum of Natural History.

ANSP Academy of Natural Sciences of Philadelphia.

CNHM Chicago Natural History Museum.

EHT-HMS Personal collection of Dr. Edward H. Taylor.

IBM Instituto de Biologia de Mexico.

MCZ Museum of Comparative Zoology.

RTM Personal collection of R. T. Moore.

TAM Texas A & M College.

UMMZ University of Michigan Museum of Zoology.

USNM United States National Museum.

* Amer. Mid. Nat. 1949 in press.

Tihen: Lizard Genus Barisia 219

Barisia Gray

Barisia Gray, 1838, Ann. Mag. Nat. Hist., ser. 1, 1:390.

Tropidogerrhonotus Fitzinger, 1843, Syst. R'ept. :21. (genotype — rudicollis)

Barissia Gray, 1845, Cat. Spec. Liz. Coll. Br. Mus. :54.

Tropidogerrhum Agassiz, 1846, Nomencl. Zool., Ind. Univ. :203. (genotype — rudicollis)

Mesaspis Cope, 1877, Proc. Anier. Philos. Soc, 17 :96. (genotype — moreleti)

Genotype: Gerrhonotus imbricatus Wiegmann, 1828 [= Barisia
imbricata imbricata (Wiegmann) ] .

Definition: Gerrhonotine lizards with the skull not widened or
depressed ; frontal bone not in contact with the maxillae ; pterygoid
teeth absent or vestigial; dorsal osteoderms with a well-defined,
strongly thickened basal area; lateral fold moderately to well
developed, with a moderate or large granular area ; sides of the neck
finely granular; postero-ventral surfaces of the forearms and shanks
granular; minimum number of scales in a single row across the nape
six to ten; anterior internasals present; postrostral present or ab-
sent; subocular and postocular series well differentiated from each
other; the suboculars extend to the lowest primary temporal; twelve
to fourteen ventral scale rows.

Range: Panama northward through Central America and Mexico
to Chihuahua and Coahuila.

A full description of the genus may be found in my previous paper
referred to above ; it will not be repeated here.

There are three major lines of specialization within the genus;
these have given rise to three species groups, which may be defined
very briefiy as follows:

1. Postmental unpaired ; superciliaiy series usually

complete moreleti group

2. Postmental paired; superciliary series complete. . .gadovii group

3. Postmental paired; superciliary series

incomplete imbricata group

A Key to the Forms of the Genus Barisia

1. Postmental unpaired (nasal bones in contact) 2

Postmental paired ; (nasal bones separated by fronto-premaxillary contact) 8

2. Eighteen to twenty longitudinal dorsal scale rows 3

Fourteen to sixteen longitudinal dorsal scale rows 7

3. Upper and lower postnasals in contact 4

Upper postnasal separated from lower by a loreo-nasal contact moreleti rafaeli

4. Lowest primary temporal usually in contact with only the lowest secondary; pre-

frontals usually absent ; posterior loreal usually not in contact with the

supralabials 5

Lowest primary temporal usually in contact with the two lower secondaries ; pre-
frontals usually present ; posterior loreal normally in contact with the
supralabials 6

5. Anterior superciliary element frequently in contact with prefrontal; frontonasal

usually touching frontal; 52-58 dorsal scale rows '. moreleti temporalis

220 The University Science Bulletin

Anterior superciliary element not in contact with prefrontal; frontonasal usually

separated from frontal by prefrontals; 47-55 dorsal scale rows

moreleti salvadorensis

6. Anterior sublabial usually in contact with the second infralabial ; general color

frequently brownish vioreleti fulva

Anterior sublabial usually in contact with only the third, or more posterior, infra-
labial ; general color usually olive to blackish moreleti moreleti

7. Usually 16 dorsal scale rows; frontonasal present, large monticola

Fourteen dorsal scale rows ; frontonasal normally absent viridifla va

8. Superciliary series complete ; (pterygoid teeth absent) 9

Superciliary series incomplete; (pterygoid teeth vestigial) 12

9. Supranasals unexpanded; an anterior canthal present 10

Supranasals expanded ; no anterior canthal 11

10. Scales of neck keeled; anterior loreal in contact with anterior canthal. .gadovii gadovii
Scales of neck smooth ; anterior loreal separated from anterior canthal by a
contact of the posterior loreal with the upper postnasal gadovii levigata

] 1 . Frontonasal and postrostral present modesta

No frontonasal or postrostral antauges

12. One loreal ; more than 33 dorsal scale rows 13

Two loreals; fewer than 33 dorsal scale rows rudicollis

13. More than one (usually three) superciliaries ; transverse dorsal scale rows fewer

than 46 14

A single (middle) superciliary element; transverse dorsal scale rows 46 or more. . .

levicollis

14. Fewer than 16 longitudinal dorsal scale rows; contact of the anterior superciliary

with the loreal usually smaller than that of the first medial supraocular with

the loreal ivibricata imbricata

Sixteen longitudinal dorsal scale rows ; contact of the anterior superciliary with
the loreal as great or greater than that of the first medial supraocular with
the loreal 15

15. Lowe.st primary temporal in contact with the penultimate as well as the ante-

penultimate supralabial ; 39-45 transverse dorsal scale rows imbricata ciliaris

Lowest primary temporal not in contact with the penultimate supralabial; 35-39
transverse dorsal scale rows imbricata planifrona

The moreleti group

Definition: Barisia with an unpaired postmental; nasal bones
in contact with each other in the mid-ciorsal hne; pterygoid teeth
absent; small lizards, the snout-vent length never exceeding 95 mm.,
usually much less ; dorsal color dark olive brown or brown to black-
ish, with a more or less well-defined dorsal longitudinal band;
venter usually spotted or mottled with dark; superciliary series
usually complete; upper postnasal present; loreo-canthal series
variable; frontonasal and postrostral present or absent; lateral fold
moderately developed.

Range: Panamca northward to Oaxaca.

Discussion: The possession of an unpaired postmental is a uni-
versal characteristic of the group. The nasal bones are known to
be in contact in viridijiava and m. moreleti; this is assumed to be
the condition in the other members. Some examples of all species
(but not of all subspecies) have been examined to determine the
condition of the pterygoid, and in all cases the pterygoid teeth were
completely lacking.

Tihen: Lizard Genus Barisia 221

These are the smallest of all the Gerrhonotinae ; the largest species
of the group is moreleti itself, with a maximum observed snout-
vent length of 92 millimeters. B. viridiflava is the smallest form,
with an observed maximum of 71 mm. The color pattern, though
varying in detail, is basically similar in all members of the group.
A similar type of pattern is found in the gadovii group, but the
imbricata group is, in general, much lighter, with a strong tendency
towards obliteration of the dorsal longitudinal band.

The arrangement of the lateral supraoculars and the superciliary
series merits some discussion. In the majority of specimens of
moreleti there are three lateral supraoculars and a normal, complete
superciliary series. In viridiflava and monticola the usual condition
appears to be the possession of only two lateral supraoculars, with
the superciliary series complete and the most posterior element
occasionally somewhat enlarged. In some specimens of moreleti
and monticola an arrangement is found which appears very similar,
but which seems better interpreted as a loss of the posterior part
of the superciliary series, the third lateral supraocular dropping
down to contact the orbit. A very similar condition is found
throughout the imbricata group. Neither interpretation — ^the loss
of the posterior superciliaries or the loss of the third lateral supra-
ocular with an enlargement of the most posterior superciliary —
adequately accounts for all the conditions found. I have used the
latter interpretation to explain the arrangement as found in viridi-
flava, the former to explain the slightly different one found occa-
sionally in other members of this group and universally in the im-
bricata group. Possibly neither is entirely correct.

In the present group, the upper postnasal is always present,
although not necessarily in contact with the lower. The supranasal
is variable, usually present and unexpanded, but sometimes absent
in certain forms, and occasionally expanded in others. The loreo-
canthal group is quite variable in moreleti itself but usually con-
taining at least three elements, while in other species, only one or
two elements are usually present.

The group as a whole shows a strong tendency towards enlarge-
ment of the frontonasal, with corresponding reduction of the pos-
terior internasals and, more particularly, the prefrontals; these
latter are in some cases completely lacking, either dropped out
or fused with the enlarged frontonasal. In one form, viridiflava,
modification has been in*the opposite direction; the frontonasal is
absent or, in those exceptional individuals where it does occur, is

222 The University Science Bulletin

relatively small. The postrostral is normally lacking in all forms,
but appears in occasional specimens of viridiflava.

The dorsal osteoderms are rather thin, slightly rugose, with a
strongly thickened basal portion and a rather weak and obtuse
median keel. The lateral fold is only moderately developed, the
included granular area being about equivalent to one lateral scale
row in width. The side of the neck contains a very poorly-defined
dermal pocket or none at all.

The moreleti group, particularly the species moreleti, is thought
to be the most primitive of the genus. The unpaired postmental,
reduction of the prefrontals, and complete lack of pterygoid teeth
are modifications not found in the primitive Barisia stock, but in
totality of characters this group approaches that stock more closely
than does any other.

I do not at present propose to enter into great detail regarding
subspecific differentiation within the species Barisia moreleti. There
is much individual variation in this form, and the localities from
which collections of series of individuals have been made are so
scattered that areas of intergradation cannot yet be definitely estab-
lished. The taxonomic arrangement is therefore far from settled,
and I prefer to give at present only a brief resume of those forms
which I believe to be valid subspecies.

Barisia moreleti moreleti (Bocourt)

Gerrhonotus moreleti Bocouvt, 1871, Nouv. Arch. Mus., 7, Bull. :102.
Gerrhonotus moreletii, Bocourt, 1871, ibid. :103.
Mesas-pis inoreletii, Cope, 1877, Proc. Amer. Philos. Soc, 17 :96.

Gerrhonotus [moreletii] m,oreletii, Dunn and Emlen, 1932, Proc. Acad. Nat. Sci. Philad.,
84:29.

Type and type locality: A series of specimens presumably in
the Paris Museum. From "le Peten, ainsi que les forets de pins de
la Haute Vera-Paz (Guatemala)."

Range: Alta Verapaz, Guatemala, westward possibly to, but not
beyond, the Cuchumatanes, and southward into Honduras, where
presumed intergrades with salvadorensis are found.

Diagnosis: A Barisia of the moreleti group with 18-20 longitu-
dinal rows of dorsals; transverse rows of dorsals 49-56; lowest pri-
mary temporal in contact with only the lowest secondary; supra-
nasals present, unexpanded; frontonasal present, large, usually in
contact with the frontal; prefrontals usually absent; upper and
lower postnasals in contact with each ot^er; anterior superciliary
not in contact with prefrontal (when present) ; posterior loreal
usually separated from the supralabials; sublabial series usually

Tihen: Lizard Genus Barisia 223

extending anteriorly only to the third or a more posterior infra-
labial ; dorsal color usually dark brown or olive brown to blackish ;
venter usually relatively dark in color, with a rather conspicuous
light V-shaped marking on the chin.

Barisia moreleti fulva (Bocourt)

Gerrhonotus fulvus Bocourt, 1871, Nouv. Arch. Mus., 7, Bull. :104.

Mesaspis fulvus, Cope, 1877, Proc. Amer. Philos. Soc, 17:96.

Gerrhonotus moreletii fulvus, Stuart, 1943, Gee. Pap. Univ. Mich. Mus. Zool., No. 471:20.

Type and type locality: A series in the Paris museum; from
"les forets de pins de Totonicapan (Guatemala)."

Range: Northwestern Guatemala, the limits of the range not
established. Specimens from the Sierra de los Cuchumatanes are
not fully typical of this form; they may possibly represent three-
way intergradation — moreleti, m. fulva and m. temporalis, but they
also present certain unique characteristics, indicating that the situ-
ation may be even more complex.

Diagnosis: Similar to B. m. moreleti, except: transverse rows of
dorsals 50-59 ; sublabial series usually extending anteriorly to touch
the second infralabial; color generally somewhat lighter, more
brownish dorsally; venter also somewhat lighter, therefore the V-
shaped marking on the chin less conspicuous.

Note: If the locality data concerning the type are correctly given
by Bocourt, the name julva seems the most likely name for this
western Guatemala subspecies. However, the figure of this form
subsequently published by him in his "Mission Scientifique," PI.
21b, figs. 6, 6a, shows certain features definitely not characteristic
of the population to which the name is here assigned. Since these
features were not mentioned in the original description, further
study is required to determine whether or not the figure actually
represents one of the type series of julva, and if so, whether it is
only an aberrant individual or is typical of that series, and whether
or not the locality data are correct, before a certain assignation of
the name can be made. Since this certain assignation cannot be
made at present, I believe it will cause the least confusion to con-
tinue, for the present, to apply this name to the western Guatemala
form.

Barisia moreleti rafaeli (Hartweg and Tihen)

Gerrhonotus vioreleti rafaeli Hartweg and Tihen, 1946, Gee. Pap. Univ. Mich. Mus. Zool.,
No. 497:8.

Type and type locality: UMMZ 88228; a young female from
16 km. S of Siltepec, Chiapas.

224 The University Science Bulletin

Range: The southern Sierra de Chiapas. Known from the fol-
lowing localities: 16 km. S of Siltepec, Chiapas (UMMZ 88227-
88228), Cerro Paxtal (UMMZ 88384), Chiquihuite, Volcan de
Tacana (UMMZ 88384), Cerro Male (UMMZ 94290), and Volcan
Tajiimiilco, Guatemala (CNHM 20308-20310).

Diagnosis: Similar to B. m. moreleti except: dorsals in 50-55
transverse rows; prefrontals present; upper and lower postnasals
separated from each other by the anterior loreal, which is in con-
tact with the nasal; posterior loreal in contact with the suprala-
bials; V-shaped marking on chin less distinct, the marking itself
being darker than in m. moreleti.

Barisia moreleti salvadorensis (Schmidt)

Gerrhonotus salvadorensis Schmidt, 1928, Field Mus. Nat. Hist. Zool. Ser., vol. 12, nc
16, 196.

Gerrhonotus moreletii salvadorensis, Dunn and Emlen, 1932, Proc. Acad. Nat. Sci. Philac
84, p. 28.

Type and type locality: CNHM 10957; an adult female .:im
Los Esesmiles, dept. Chalatenango, Salvador.

Range: Honduras and Salvador southward to Matagalpi, Nica-
ragua.

Diagnosis: Similar to B. m. moreleti, except: transverse rows of
dorsals 47-55 ; lowest primary temporal usually in contact with the
two lowest secondaries; frontonasal usually separated from the
frontal by the prefrontals, which are consistently present; posterior
loreal in contact with the supralabials; sublabials extending an-
teriorly to touch the second infralabials.

Barisia moreleti temporalis (Hartweg and Tihen)

Gerrhonotus moreleti temporalis Hartweg and Tihen, 1946, Dec. Pap. Univ. Mich. Mus.
Zool., No. 497:10.

Type and type locality: UMMZ 94910; an adult male, collected
11 km. southeast of Ciudad de las Casas, Chiapas.

Range: Known only from the vicinity of Ciudad de las Casas,
Chiapas.

Diagnosis: Similar to B. m. moreleti, except: 52-58 transverse
rows of dorsals; lowest primary temporal usually in contact with
the two lowest secondaries; prefrontals present, sometimes sepa-
rating frontonasal from frontal; anterior superciliary frequently in
contact with prefrontal; posterior loreal in contact with the supra-
labials; sublabials extending anteriorly to the second infralabial;
dorsal and ventral coloration both decidedly lighter in tone, the

Tihen: Lizard Genus Barisia 225

V-shaped chin marking less conspicuous because of the lighter back-
ground.

General remarks: The species moreleti is the most generalized
member of the group, and of the genus Barisia. This is demon-
strated by the high number of dorsal scale rows, the possession of
three lateral supraoculars and a complete superciliary series, the
relatively large number of loreo-canthal elements, and the general
coloration. There are, of course, certain features considered as
highly modified, particularly the reduction or loss of the prefrontals
and, in common with the other members of the group, the complete
lack of pterygoid teeth and the possession of a single postmental.

Barisia monticola (Cope)

Plate II, figs. 1, 2

Gerrhonotus monticolus Cope, 1877, Proc. Amer. Philos. Soc, 17:97.

Gerrhonotxis alfaroi Stejneger, 1907, Proc. U. S. Nat. Mus., 32 no. 1542:505. (Type
locality: Santa Maria [de Dota], Costa Rica.)

Type and type locality: In the collection of the U. S. National
Museum; number not designated by Cope. From the summit of
Pico Blanco in Costa Rica.

Range: The mountains of western Costa Rica south to Chiriqui,
Panama.

Diagnosis: A Barisia of the moreleti group with sixteen longi-
tudinal rows of dorsals (the most lateral occasionally much re-
duced) ; transverse rows of dorsals 43-51; supranasals present or
absent; frontonasal present; prefrontals present, often much re-
duced, or occasionally absent.

Description: No postrostral. Nasal normally separated from
rostral by the anterior internasals. Supranasals usually absent.
Frontonasal present, large, in contact wuth the frontal or not; pre-
frontals usually present, often much reduced. Loreo-canthal region
variable, but seldom more than two elements present. One pre-
ocular; two or three suboculars; two to four postoculars. Three to
five superciliaries, the series complete posteriorly, with the posterior
element frequently much enlarged; two, sometimes three, lateral
supraoculars. Four primary and four secondary temporals, the
uppermost primary in contact with the uppermost secondary or
not, and the lowest primary normally in contact with only the low-
est secondary, occasionally the two lowest. Supralabials 8-10; in-
fralabials 7-9. Postmental unpaired, followed by three large pairs
and one smaller pair of chin shields, of which the members of the
first, occasionally also the second, pair are in contact along the mid-
15— 3659

226 The University Science Bulletin

ventral line. The sublabials extend anteriorly to the first chin
shields (rarely the postmental) and the second infralabials.

Dorsals in sixteen longitudinal rows, the most lateral occasionally
reduced, and 43-51 (ave. 47.7) transverse rows. Ventrals in twelve
longitudinal and 51-57 (ave. 53.9) transverse rows. Caudal whorls
78-87 (ave. 83.1). The median six or eight rows of dorsals are
rather weakly and obtusely keeled.

There is a marked sexual dimorphism in the color pattern. Males
black or brownish-black dorsally, with many small bluish-white
spots (green in life), these spots somewhat larger and more distinct
dorsally than laterally, producing a poorly defined dorsal longi-
tudinal band. A latero-dorsal dark line bordering this band, and
an interrupted mid-dorsal dark stripe. Ventral surface mottled
(yellow green in life), the chin and throat frequently lighter than
the belly; a more or less distinct V-shaped mark along the chin
shields. Females a much lighter brown, with the whitish spots less
numerous or lacking. The dorso-lateral dark line is very promi-
nent, often bordered above by a narrow light line. A similarly
prominent interrupted mid-dorsal stripe. Belly much lighter than
in males, tending towards a uniform grayish and not prominently
mottled.

These are small lizards, averaging around 78 mm. in snout-vent
length; the observed maximum is 87 mm. Tail 1.58-1.68 times the
snout-vent length.

Remarks: This species shows certain resemblances to B. morcleti,
but differs from that species in the smaller number of dorsal scale
rows, both transverse and longitudinal, and in color pattern, par-
ticularly of the males. It also differs in the frequent absence of
supranasals. Although the loreo-canthal region is somewhat var-
iable within both species, there is a definite tendency for monticola
to possess a smaller number of elements in this region. The num-
ber of supralabials is, in monticola, most frequently nine, in more-
leti most frequently ten. In view of the lack of intergrading speci-
mens and the fairly numerous and constant differences between
monticola and all races of m,oreleti, it seems best for the present to
retain full specific rank for monticola.

Certain variable features within this species give indications that
they may possess geographic significance. The presence or absence
of supranasals and the contact or lack of a contact of the upper-
most primary with the uppermost secondary temporal may also
prove to have geographic significance. Material at present avail-

Tihen: Lizard Genus Barisia 227

able, is however, inadequate to conclusively demonstrate the ex-
istence of more than one race.

Specimens examined: Costa Rica: Barba (MCZ 28077-28078); Irazii (MCZ 15465,
15468, 32079); Poas (MCZ 15466-15467); Cerro de las Vueltas (USNM 70649); Volcan
Irazii (20) 8,000-10,000 ft.; Volcan Poas (3) 6,300 ft. Cerro de la Muerte (1) 11,000 ft.
(uncatalogued specimens). Specimens have also been reported from Pico Blanco and Santa
Maria, Costa Rica. Panama: Chiriqui Volcano (MCZ 45664; USNM 94991).

Barisia viridifiava (Bocourt)

Figure 1

Gerrhonotus viridiflavus Bocourt, 1873, Ann. Sci. Nat., ser. 5, 17, art. 2 (unpaged).

Gerrhonotus bocourti Peters, 1877, Monatsber. K. Preus. Akad. Wiss. Berlin, 1876 :297.
(Type locality: "Mexico.")

Barissia bocourti, Cope, 1877, Proc. Amer. Philos. Soc, 17 :97.

Gerrhonotus obscunis G'iinther, 1885, Biol. Centr. Amer., Rept. : 40. (Type locality:
"Mexico.")

Type and type locality: In the Paris Museum, number unknown.
The locality is given merely as ''Mexique," but in view of the pres-
ent known distribution, I propose to restrict the type locality to
"the highlands of central Oaxaca."

Range: Highlands of central Oaxaca.

Diagnosis: A Barisia of the moreleti group with fourteen longi-
tudinal rows of dorsals; transverse rows of dorsals 48-54; supra-

FiG. 1. Barisia viridifiava U. S. National Museum no. 113220. Actual
snout to ear measurement — 13.8 mm.

228 The University Science Bulletin

nasals present, expanded or not; frontonasal normally lacking; pre-
frontals present, not reduced in size.

Description: Postrostral present or absent; occasionally two an-
terior internasal elements present on each side. Supranasals pres-
ent, occasionally expanded to meet the postrostral (when present)
or, more rarely, to meet each other in the mid-dorsal line. Fronto-
nasal normally absent, rarely present. One loreal; no canthals.
The usual arrangement of the scales above the eye is interpreted
as follows: two lateral supraoculars; the superciliary series is com-
plete, with the most posterior element usually somewhat enlarged.
Three to five superciliaries, the most anterior only rarely in contact
with the prefrontal. Suboculars two; postoculars three. Normally
four primary and four secondary temporals, the uppermost primary
occasionally in contact with the uppermost secondary, more fre-
quently not, and the lowest primary in contact with only the lowest
secondary. Nine, occasionally eight, supralabials; seven or eight
infralabials. Postmental unpaired, followed by three large pairs
and one much smaller pair of chin shields, of which the members
of the anterior pair are in contact mid-ventrally ; the members of
the second pair are frequently in contact with each other posterior
to a single small median scale intercalated between these and the
first pair. The sublabials extend anteriorly to the first chin shield
and second infralabial.

Dorsals in fourteen longitudinal and 48-55 (av. 51.6) transverse
rows; ventrals in twelve longitudinal and 55-61 (av. 57.9) trans-
verse rows. Caudal whorls 83 and 87 in the only two specimens
seen with undamaged tails. The median six or eight rows of dorsals
are rather weakly keeled.

Color pattern similar in general to that of B. m. moreleti, but with
great variation in details. The dorsal longitudinal band may be
veiy prominent or very poorly defined; the sides may show very
many or practically no white-tipped scales; the belly is usually
very dark, but occasionally a very light gray, and may be strongly
mottled or nearly uniform. These variations do not seem to be
correlated with age, sex, or locality, excepting that the Mt. Zem-
poaltepec specimens have, on the whole, a better defined pattern,
with stronger contrasts between the various elements, than do speci-
mens from other localities.

This is the smallest of all the gerrhonotine lizards, adults aver-
aging slightly over 60 mm. in snout- vent length, with an observed
maximum of 71 mm. Tail 1.55-1.67 times the snout-vent length.

Tihen: Lizard Genus Barisia 229

Remarks: Since the characters presumably distinguishing the
three forms, viridifiava, bocourti and obscurus are all found in a
single series of specimens from a single locality (Cerro San Felipe),
and since specimens from other localities show no evidence of any
geographic significance in these variations, I consider the three
names synonymous, with viridiflava having priority.

The species is certainly a member of the moreleti group, but
differs rather widely from the other members, and is considered to
be the most highly modified form in the group. The low number
of longitudinal dorsal scale rows, the constant absence of all but
one loreo-canthal element, the small size, the usual absence of the
frontonasal, the usual occurrence of only two lateral supraoculars,
and the relatively low number of supralabials are all considered
specialized characters, and are the ones which serve to distinguish
this form from the other members of the group.

Specimens examined: Oaxaca. Mt. Zempoaltepec (RTM 394, 403-404, 406; USNM
47184-47185, 47599); Cerro San Felipe (EHT-HMS 19146-19149, 23810-23835); Chivato
(EHT-HMS 28133-28134); Cuicatlan (USNM 47612). A total of forty specimens; no other
specimens with definite locality data have been reported.

The gadovii group

Definition: Barisia with a paired postmental; nasal bones sep-
arated from each other by a narrow contact of the frontal with the
premaxilla; lizards of small to moderate size, not exceeding 110
mm. in snout-vent length; coloration similar to that of the moreleti
group, but with a greater tendency for vertical bars along the sides,
occasionally continued across the back; superciliary series com-
plete; upper postnasal present; usually (?) two loreals; an anterior
canthal may or may not be present; frontonasal and postrostral
present or not; lateral fold moderately developed.

Range: Southern Guerrero through Oaxaca to Veracruz.

Discussion: I have chosen to term this group the gadovii group,
rather than assign to it the older name of antauges, because the
latter form is so poorly known that its affinities are uncertain. It
has been tentatively referred to this group on the basis of its pos-
session of a paired postmental; the same is also true of B. modesta.
Since these forms are so inadequately known, the group has been
defined chiefly on the basis of the species gadovii. Further inform-
ation concerning these other forms may require a modification of
this definition, or may show that they have been incorrectly assigned
to this group.

The gadovii group is in many respects intermediate between the

230 The University Science Bulletin

moreleti and imbricata groups, morphologically as well as geo-
graphically. It possesses the relatively liberal scalation and general
color pattern of the former, and also resembles that group in the
relatively weak development of the lateral fold and the complete
lack of pterygoid teeth. It agrees with the imbricata group in the
possession of a paired postmental and in the important osteological
feature of the separation of the nasals by a fronto-premaxillary
contact, though this separation is narrower here than in the im-
bricata group. In size and color pattern it is also more or less in-
termediate between the other two, but closer to the moreleti group.

Barisia gadovii gadovii (Boulenger)

Gerrhonotus gadovii Boulenger, 1913, Ann. Mag. Nat. Hist., ser. 8, 12 :564.

Type and type locality: A series of specimens in the British
Museum, no holotype designated. This series is from Omilteme,
Guerrero.

Range: Known only from the vicinity of Omilteme, Guerrero,
but possibly ranging southward into Oaxaca.

Diagnosis: A Barisia of the gadovii group with a frontonasal;
supranasals small, unexpanded; lateral scales as well as dorsal scales
of body keeled; those of neck also with keels; usually two loreals
and one (anterior) canthal, the latter in contact with the anterior
loreal; belly spotted or mottled with black; dorsal color a dark
brownish gray or brownish black.

Description: Postrostral present or absent. Supranasals present,
small, unexpanded. Frontonasal present, in contact with the an-
terior canthal and often with the frontal. Two loreals and one
(anterior) canthal, the latter normally in contact with the anterior
loreal. Normally three lateral supraoculars. Superciliaries three
to six, the most anterior not in contact with the prefrontal. Usually
one preocular, one subocular, and two or three postoculars. Four
primary and four secondary temporals, the uppermost primary
usually in contact with the uppermost secondary, and the lowest pri-
mary with only the lowest, occasionally also the second, secondary.
Supralabials 8-10; infralabials 7-9. Postmental paired, followed
by three large pairs and one smaller pair of chin shields, of which
the members of the first pair, frequently also the second, are in
contact along the mid-ventral line. The sublabials extend anteri-
orly to the first chin shields and second infralabials, occasionally
the postmental and/or first infralabial.

Dorsals in 16-18 longitudinal and 46-51 (ave. 48.4) transverse

Tihen: Lizard Genus Barisia 231

rows; ventrals in twelve longitudinal and 52-56 (ave. 53.9) trans-
verse series. Occasionally some of the transverse rows of ventrals
contain only ten or eleven scales. Caudal whorls 78-88 (ave. 84.2).
Usually all or nearly all of the dorsal scales are prominently keeled,
as are those of the sides of the tail and one or two rows on the thigh
and on the shank.

A dark brownish gray or brownish black dorsally, often with a
more or less distinct longitudinal band. Narrow vertical black bars
on the sides, usually bordered posteriorly with white; these bars
may be continued across the back as obtusely V-shaped markings.
There are about twelve such bars between the axilla and the groin.
Venter heavily spotted or mottled with black in males, lightly in
females. A dark line from the nasal across the eye onto the tem-
porals, bordered below by a very prominent light stripe.

This form is of moderate size, adults averaging about 90 mm. in
snout-vent length, the observed maximum being 102 mm. Tail
1.44-1.72 (ave. 1.59) times the snout-vent length.

Remarks: The species gadovii does not appear to be particularly
closely related to any other. Oaxacan specimens represent a dif-
ferent form, but are thought to constitute a subspecies of gadovii
rather than a distinct species; differences are discussed in connec-
tion with that form.

There is a slight superficial resemblance between this form and
Elgaria multicarinata of the United States Pacific Coast. Some
multicarinata reported from "Mexico" may well be referable to this
form. In this connection, there can be little doubt that the figure
of a skull in Bocourt's "Mission Scientifique" (PI. 21c, fig. 7, 7a)
and labelled "multicarinata" actually represents a specimen of
gadovii.

Specimens examined: Guerrero: Omilteme and Chilpancingo (USNM 47737, 113173-
113174; MCZ 42701-42715; BHT-HMS 23849-23876; CNHM 38516-38524). A total of
fifty-six specimens. The form is not known from any other locality.

Barisia gadovii levigata, subsp. nov.

Plate II, figs. 3, 4, 5.

Type and type locality: Holotype: USNM 47212; an adult male
from the "Valley of Oaxaca." Paratype: USNM 47855; an adult
male from the "mountains west of Oaxaca City." Nelson and Gold-
man colls.

Range: Known only from the localities mentioned above.
Diagnosis: A Barisia of the gadovii group with a frontonasal;
supranasals small, unexpanded; scales of the sides of the body

232 The University Science Bulletin

smooth, those of the back weakly keeled, and those of the neck
smooth; two loreals and one (anterior) canthal, the latter separated
from the anterior loreal by a contact of the posterior loreal with
the upper postnasal; belly spotted with brown; dorsal color a light
brownish gray.

Description of the holotype: No postrostral; nasal separated
from the rostral by the anterior internasals. Siipranasals small,
imexpanded. Frontonasal in narrow contact with the frontal be-
tween the prefrontals. Two superposed postnasals. Two loreals
and one (anterior) canthal, the latter narrowly separated from the
anterior loreal by a contact of the posterior loreal with the upper
postnasal. One preocular; one subocular; four postoculars, the low-
est quite small. Superciliary series incomplete posteriorly, com-
prising four elements on the left, only two on the right — the three
anterior being fused into a single long element; the most anterior
superciliary is not in contact with the prefrontal. Five medial and
three lateral supraoculars, the posterior lateral in contact with the
orbit. Frontal touching the interparietal. Four primary and four
secondary temporals, the uppermost primary in contact with the
uppermost secondaiy, and the lowest primary in contact with only
the lowest secondary. Four temporal elements in contact with the
supralabials. Supralabials nine on the left, ten on the right; infra-
labials eight. Postmental paired, followed by three large and one
much smaller chin shields on each side, of which the members of
the first pair are in contact along the mid-ventral line. Four or five
sublabials, the anterior extending to the first chin shield and second
infralabial.

Dorsals in sixteen longitudinal and forty-seven transverse rows;
ventrals in twelve longitudinal and fifty-one transverse rows. A
minimum of eight scales in any single row across the nape. The
lateral rows of dorsals are smooth, the median eight rows weakly
keeled except on the neck, where all of the scales are smooth. All
scales of the limbs and those of the sides of the tail smooth or very
weakly keeled.

A light brownish gray dorsally, each individual scale finely
mottled brown and gray. Sides with somewhat irregular blackish
vertical bars, bordered posteriorly with white; about 10-12 such
bars between the axilla and the groin, the more posterior ones in-
distinct. A dark line from the nasal through the lower part of the
eye onto the lower temporal region, bordered below by a very promi-
nent light stripe. Ventral surface a very light gray or yellowish,

Tihen: Lizard Genus Barisia 233

with many brown spots which tend strongly to be arranged in longi-
tudinal lines along the middles of the scale rows, particularly me-
dially. Chin and infralabials with numerous irregular small brown
dots, the infralabial region not banded.

Measurements of the holotype are as follows:

Snout-vent 107 mm. Axilla-groin 55 mm.

Tail regenerated Fore limb 24 mm.

Head length 24.4 mm. Hind limb 30 mm.

Head width 16.8 mm. Fourth finger 7.1mm.

Head depth 13.5 mm. Fourth toe 11.3 mm.

Description of the paratype: This specimen agrees with the holo-
type except as here noted.

Frontonasal separated from the frontal by the prefrontals. Three
postoculars. Superciliary series complete, comprising three ele-
ments on the left, four on the right; four lateral supraoculars on
the left, three on the right, none of which are in contact with the
orbit. Probably nine supralabials on each side, but because of an
injury this number is not entirely certain. Seven infralabials.

Transverse rows of dorsals forty-five; ventrals in fifty-two trans-
verse series.

Somewhat lighter in color than the holotype; each dorsal scale
primarily gray, very finely dotted with brown.

Measurements of the paratype are as follows:

Snout-vent 97 mm. Axilla-groin 47 mm.

Tail 130 mm. Fore limb 21 mm.

Head length 21 .2 mm. Hind limb 27 mm.

Head width 15.3 mm. Fourth finger 6.0 mm.

Head depth 12.8 mm. Fourth toe 9.7 mm.

Remarks: This form is closely related to B. g. gadovii, from
Vvhich it differs chiefly in the separation of the anterior loreal from
the anterior canthal, the much less extensive carination, and the
lighter color, with brown rather than black ventral spots ; probably
also in the possession of a slightly lower number of transverse
dorsal scale rows. The difference in coloration might conceivably
be due to differences in manner of preservation, but this seems veiy
unlikely in view of the large number of g. gadovii seen, comprising
series from several different collections; included in these are Guer-
rero specimens collected by Nelson and Goldman, the collectors of
the types of levigata.

The close general resemblance between this form and typical
gadovii makes a subspecific relationship appear more probable than
a specific one.

Specimens examined: Oaxaca: Valley of Oaxaca (USNM 47212, holotype); "Mountains
west of Oaxaca City" (USNM 47855, paratype). No other specimens are known.

234 The University Science Bulletin

Barisia modesta (Cope)

Pterogasterus modestus Cop€, 1877, Proc. Amer. Philos. Soc, 17 :97.
Gerrhonotus modestus, Gunther, 1885, Biol. Centr. Amer., Rept. : 42.

Type and type locality: USNM 7084 (three specimens). Tlie
type locality is uncertain. Cope states, "The precise locality from
which the specimens of this lizard were sent to the Smithsonian
Institution is uncertain, but is probably Guatemala." Dr. Hobart
Smith informs me that the tags on these specimens bear the datum
"Veracruz," and has stated (1942) that the "type locality [is] ap-
parently Orizaba, not Guatemala as guessed by Cope."

Range: Uncertain; the form is known only from the types.

Diagnosis: A Barisia of the gadovii group with frontonasal;
supranasals expanded; dorsal scales of the body and neck nearly
smooth; two (?) loreals, no canthals (?) ; dorsal color "brown . , .
the sides are a reddish brown or maroon . . ."

Description: I have not seen this form, and all descriptions to
be found in the literature are derived from Cope's original descrip-
tion, which is copied below. The bracketed comments are mine.

"Scales 10/12 [a misprint for 16/12, c/. Smith, 1942] slightly
convex above, but not keeled, excepting those of the tail, which are
strongly and obtusely carinate or ribbed; an azygous scute [post-
rostral] between the two anterior pairs of internasals [anterior
intemasals and expanded supranasals]. Internasals of first pair
reaching first labials. Internasals of third pair [posterior inter-
nasals] elongate, in contact with the frontonasals [prefrontals] be-
hind, apparently including the small lateral interfrontonasals [an-
terior canthals?]. Two postnasals; a large plate, the anterior [pos-
terior?] canthal, descends to the labials, from the inferior part of
which a loreal may be separated. [I have not been able, from this
description, to satisfy myself as to the exact arrangement of scales
in this region.] Preoculars two or one. Two pairs of large infra-
labials [postmentals and anterior chin shields] in contact, follow-
ing the symphyseal, without a single postmental; two pairs follow,
of which the anterior are separated by one scute. Lateral fold ex-
tending from ear to vent; the granular area extending above the
humerus. Appressed limbs separated by six cross-rows of ab-
dominal scales, or the length of the fore arm. Rows of scales from
nape to origin of tail, forty-seven ; do. from humerus to vent, thirty-
eight.

"The tail is not very long and is grooved below as well as above.
Total length, .150 [presumably meters, hence 150 mm.] ; length to

Tihen: Lizard Genus Barisia 235

meatus auditorius, .012; to vent, .072; length of hind leg, .019. Color
above, brown; below, olivaceous. The sides are a reddish-brown or
maroon, bordered above by a blackish line which separates it from
the dorsal color.

"This species differs from all others of the genus 'Pterogasterus'
in the extinction of the small plate [anterior canthal] which trun-
cates the lateral angle of the interfrontonasal [frontonasal]. As a
consequence of this, the latter has a diamond shape, as it does not
reach the frontal plate behind nor the azygous plate in front. The
smooth scales also separate it from all others of the genus.

"The precise locality from which the specimens of this lizard were
sent to the Smithsonian Institution is uncertain, but is probably
Guatemala."

Remarks: The relationships of this form are at present obscure,
but the presence of a paired postmental, a postrostral and fronto-
nasal have induced me to place it tentatively with the gadovii group.
The fact that Cope placed it in the genus Pterogasterus, in which
he otherwise included only Gerrhonotus liocephalus and related
forms, would argue a general similarity to liocephalus; such a gen-
eral similarity is also observable in B. gadovii. A better knowledge
of the scalation and information regarding the skeletal anatomy
would be of great assistance in definitely allocating this form.

SpeciTneiis examined: None; since it is known only from the types, there are no definite
locality records for this form. [At this time (1944) the types are not available for study
at the National Museum.]

Barisia antauges Cope

Barissia antauges Cope, 1866, Proc. Acad. Nat. Sci. Philad., 1866 :132.
Gerrhonotus antauges, Sumichrast, 1882, La Naturaleza, 6 :40.

Type and type locality: USNM; collected by Professor Sumi-
chrast, from Orizaba, Veracruz.

Range: Known only from the type locality.

Diagnosis: A Barisia of the gadovii group without a frontonasal;
supranasals expanded; dorsal scales of the body and neck nearly
smooth; two (?) loreals, no canthals; ventral surface very heavily
mottled; dorsal surface "dark brown, with a subdivided iridescence
as though greased."

Description: As in the case of modesta, very little is known con-
cerning this form. I have not seen the type, and all literature de-
scriptions subsequent to Cope's, but excepting the very brief char-
acterization in Smith's (1942) key, have been written by authors
who considered B. viridifiava a synonym of antauges; their descrip-

236 The University Science Bulletin

tions actually apply to the former species. Again the best pro-
cedure seems to be to copy Cope's description, and again the brack-
eted comments are mine.

"A species differing from those already known in the entire
smoothness of the scales of the body, while those of the tail are ar-
ranged in obtuse and strong ridges. Nuchal rows eight, those of
the body 16/12. A depression along the vertebral line; six scales
margin the vent. Labials 10/8, three last superior nearly equal,
separated by four rows of nearly equal temporals from the parietals.
Latter as broad as long, well separated, with the frontoparietals,
by the elongate interparietal. Five supraorbitals, embracing three
superciliaries [lateral supraoculars]. Prefrontals longer than broad;
three pairs supranasals [anterior internasals, expanded supranasals,
posterior internasals]. Head short and elevated. End muzzle to
axilla 1 in. 3 1. [approx. 32 mm.] ; latter to vent 2 in. 1 1. [approx.
53 mm., the snout-vent length thus approximating 85 mm.] ; from
latter to end of tail 4 in. 1 1. [approx. 103 mm.; it is probable that
the tail has been injured].

"Above dark brown, with a subdivided iridescence as though
greased, and with many small blackish brown spots, which are more
distinct on the tail. Sides with about seventeen irregular vertical
black bars from opposite nape to groin, each bordered with yellow
specks behind. Front of ear and lips black, yellow varied; body
and tail below, blackish, with very many yellowish-white specks.

"No. 11, Sumichrast's Coll. Stated by Prof. S. to be very rare."

Smith offers the further information that this form possesses a
paired postmental.

Remarks: As in the case of modesta, further information is neces-
sary before any definite conclusions can be drawn. The consistent
confusion of viridifiava with this form might lead one to expect a
strong general similarity between the two, but, with the exception
of Cope himself, probably none of the authors who considered the
two synonymous had seen any specimens of antauges. Smith, who
had seen the type, recognized the distinctness of viridifiava. The
paired postmental indicates a probable affinity with the gadovii
group; the size of the type specimen is also more consistent with
this group than with the smaller moreleti or larger imbricata group.

So far as can be definitely determined, antauges and modesta
differ chiefly in the presence or absence of a frontonasal and post-
rostal. If this be the case, and if modesta actually comes from
Veracruz, there is a possibility that the two forms are identical.

Specimens examined: None; known only from Orizaba, Veracruz.

Tihen: Lizard Genus Barisia 237

The imbricata group

Definition: Barisia with a paired postmental; nasal bones sep-
arated from each other by a broad contact between the frontal and
premaxilla; pterygoid teeth vestigial; moderate to large forms, the
maximum snout-vent length observed being 155 mm.; dorsal color
brown, usually decidedly lighter than in other groups, and often
without a dorsal longitudinal band; venter relatively light, some-
times suffused with gray, but not spotted or mottled over most of
the surface; superciliary series incomplete posteriorly, sometimes
anteriorly as well; no upper postnasal; usually a single (posterior)
loreal, which may have a canthal split off dorsally (a small anterior
loreal is also present in rudicollis) ; frontonasal and postrostral ab-
sent; lateral fold very well developed.

Range: Oaxaca and Veracruz northward to Chihuahua and Coa-
huila.

Discussion: In occasional specimens there is a partial fusion of
the postmental elements, but this is rarely complete; the resulting
scute is decidedly asymmetrical and obviously different from the
symmetrical unpaired postmental of the moreleti group. The fronto-
premaxillary contact is broad, much broader than in gadovii. The
pterygoid teeth are poorly developed and few in number, but close
examination shows their constant presence; only occasionally do
any protrude through the mucous lining of the roof of the mouth.

Although interspecific size differences are present, the average
adult size of every form is greater than for any member of either of
the other groups; the more northern forms are the largest. The
dorsal color is a nearly uniform brown or tan in males of all forms
excepting rudicollis. In that species, and in females of certain
races of imhncata, a dorsal longitudinal band and lateral dark bars
are present, much as in the other groups. The ventral surface tends
to be uniformly light medially, but dark markings are frequently
found laterally. There is never the heavy spotting or mottling of
the belly frequently found in the other groups.

The superciliary series is without exception incomplete posteriorly
(see discussion of this feature in connection with the moreleti
group) ; often the anterior portion is also reduced or lacking, so
that in some cases only a single element remains. Never does the
series comprise more than four elements.

The upper postnasal is lacking, the supranasal and lower post-
nasal usually being in contact at the posterodorsal angle of the
nasal. The supranasal is always present, never expanded. The

238 The University Science Bulletin

loreo-canthal group usually consists of a single large cantholoreal
element, with occasionally a canthal split off from the dorsal por-
tion; in rudicollis a small anterior loreal is also present. The fronto-
nasal is normally lacking; rarely it occurs as a very small plate
wedged between the posterior internasals and prefrontals. The
postrostral is always absent.

The dorsal osteoderms are heavy and moderately rugose. They
possess a strong, but usually obtuse, median keel and a strongly
thickened basal portion. The lateral fold is well developed, con-
taining a granular area more than equivalent to one lateral scale row
in width. The side of the neck often contains a rather well-defined
dermal pocket.

This group is the most highly modified of all Gerrhonotinae, with
the possible exception of some Abronia. Among the more impor-
tant specializations are included the consistent absence of the fron-
tonasal, consistent strong reduction of the loreo-canthal group, re-
duction of the superciliary series, loss of the upper postnasal, and
a broad fronto-premaxillary contact. Other specializations are
shared with only a few other groups.

Barisia rudicollis (Wiegmann)

Plate I

Gerrhonotus rudicollis Wiegmann, 1828, Isis, 1828, no. 3 :380.
Barisia rudicollis. Gray, 1838, Ann. Mag. Nat. Hist., ser. 1, 1:390.
Tropidogerrhonotus rudicollis, Fitzinger, 1843, Syst. Rept. : 21.
Barissia rudicollis, Gray, 1845, Cat. Spec. Liz. Coll. Br. Mus. : 55.

Type and type locality: Berlin Museum. Type locality given
only as "Mexico," but is probably Mexico (state).

Range: Except for the types the form is known only from a
single specimen, first reported by Martin del Campo (1939), from
the Hacienda La Gavia, in the state of Mexico.

Diagnosis: A Barisia of the imbricata group with fourteen rows
of ventrals; two loreals, no canthals; nasal in contact with the
rostral; fewer than 33 transverse rows of dorsals; fourteen longi-
tudinal rows of dorsals; anterior superciliary present, moderately
large.

Description: Nasal in contact with the rostral below the an-
terior internasals. Two loreals, the anterior small, the posterior
large, in contact with the prefrontal; no canthals. Postnasal in very
narrow contact with the supranasal, or separated from it by a con-
tact of the anterior loreal with the nasal, widely separated from
the posterior internasal. Four superciliaries, the anterior separat-

Tihen: Lizard Genus Barisia 239

ing the preocular from the first medial supraocular ; the first super-
ciliary large, its contact with the (posterior) loreal greater than
that of the first medial supraocular with the loreal. Two or three
suboculars; four postoculars. Four primaiy and four secondary-
temporals, the uppermost primary in contact with the uppermost
secondary, the lowest primary with the two lower secondaries and
with the penultimate and antepenultimate supralabials. Supra-
labials 10-12; infralabials 8-9. Postmental paired, followed by
three large and one or two smaller pairs of chin shields, of which
the members of the first pair are in contact along the mid-ventral
line. The sublabials extend anteriorly to the first chin shields and
second infralabials.

Dorsals in fourteen longitudinal and about 27-29 transverse rows ;
ventrals in fourteen longitudinal and about forty-seven transverse
series. All but the lateral rows of dorsals are at least lightly keeled,
the median eight rows very prominently so. The occipital, inter-
parietal and upper temporals are also keeled. Six longitudinal
rows of scales across the nape, all strongly keeled.

Dorsum a grayish brown ; sides with»prominent but very irregular
vertical black bars bordered posteriorly with white; these white
markings are continued across the back as very indistinct light
bands. Dorsum of head brownish flecked with white, lighter lat-
erally, the labial and lower temporal regions cream colored. Rostral
and each of the anterior supralabials with a narrow black stripe
along their posterior borders. A black stripe across the prcoculars
and suboculars to the antepenultimate supralabial, at the anterior
border of which it bends down sharply to the lip. Venter whitish
with irregular, prominent black markings laterally.

Measurements of the single example seen are:

Snout-vent 105 mm. Axilla-groin 51 mm.

Tail — tip broken off Fore limb 28 mm.

Head length 25.4 mm. Hind limb 34 mm.

Head width 18.5 mm. Fourth finger 8.3 mm.

Head depth 14.2 mm. Fourth toe 11.4 mm.

Remarks: Although certainly a member of the imbricata group,
this form differs widely from all others in the group. It is prob-
ably an early, but in some respects highly specialized, derivative
of the ancestral stock of this group. The fourteen rows of ventrals,
two loreals, four superciliaries and extensive carination are regarded
as primitive features indicating an early derivation. The low num-
ber of transverse dorsal scale rows, the naso-rostral contact, and
distinctive color pattern are thought to be modifications peculiar
to this form.

240 The University Science Bulletin

Nothing is known concerning the habitat of this form, but there
are some indications that it may be arboreal — in contrast to all
other members of the genus. The lichen-like color pattern is
strongly suggestive of arboreal habits. The limbs are relatively
long; only this form of all the Gerrhonotus-Barisia-Elgaria com-
plex is comparable to the arboreal Abronia in relative limb length.
The fact that a form with such a large number of primitive features
would continue to exist so near the generic center of dispersal might
be taken as an indication that this form occupies a different habitat
than its more highly modified relatives. Even the rarity of speci-
mens in museum collections lends credibility to the hypothesis that
it is arboreal. This same paucity has been characteristic of all
members of the genus Abronia until very recently, that is, until it
was discovered that they are primarily tree-dwelling forms.

Specimens examined: Mexico (state): Hacienda de la Gavia (IBM, one specimen). No
other specimens with definite locality data are known.

Barisia imbricata imbncata (Wiegmann)

Gerrhonotiis imbricatus Wiegmann, 1828, Isis, 1828, no. 3: 381.

Gerrhonotus lichenigerus Wagler, 1830, Icon. Amph. : Fasc. II, pi. 24, fig. 2. (Type
locality: Mexico.)

Gerrhonotus adspersus Wiegmann, 1834, Herp. Mex. : pi. 10. (Type locality : Mexico
[by inference].)

Barisia imbricata, Gray, 1838, Ann. Mag. Nat. Hist., ser. 1, 1: 390.

Barisia lichenigerus, Gray, 1838, ibid.: 390.

Barissia imbricata. Gray, 1845, Cat. Spec. Liz. Coll. Br. Mus. : 55.

Barissia lichenigera, Gray, 1845, ibid.: 55.

Gerrhonotus olivaceotts Baird, 1859, Proc. Acad. Nat. Sci. Philad. for 1858: 255. (Type

locality: Mexico [erroneously stated by Baird as "near San Diego, Calif."]
Barissia otivacea. Cope, 1875, Bull. U. S. Nat. Mus., 1 : 46.

Gerrhonotus [imbricatus] imbricatus, Dunn, 1936, Proc. Acad. Nat. Sci. Philad., 88: 475.

Gerrhonotus imbricatus adspersus, Smith, 1942, Proc. U. S. Nat. Mus., 92, no. 3153: 368.

Type and type locality: Berlin Museum, the type locality is
given only as "Mexico."

Range: Central Mexico (state), the Distrito Federal and More-
los, westward into Michoacan, eastward to Veracruz and southward
to central Oaxaca; northward into southern Hidalgo and probably
southern Guanajuato.

Diagnosis: A Barisia of the imbricata group with twelve rows of
ventrals; one loreal and no (occasionally one) canthals; nasal
usually separated from the rostral by the anterior internasals; 34-42
transverse and usually 14, sometimes 12, longitudinal rows of dor-
sals; dorsal color light brown, females with darker brown cross
bands, or lateral bands plus mid-dorsal blotches; coloration of ven-
ter variable; first superciliary present, moderate in size, usually
separating the preocular from the first medial supraocular; lowest

Tihen: Lizard Genus Barisia 241

primary temporal in contact with the penultimate and antepenulti-
mate supralabials.

Description: Nasal normally separated from the rostral by the
anterior internasals. One loreal, in contact with the prefrontal,
rarely with a canthal split off dorsally. Postnasal in rather broad
contact with the supranasal, widely separated from the posterior
internasal. Generally three superciliaries, the anterior usually sep-
arating the preocular from the first medial supraocular; the anterior
superciliary is of moderate to small size, its contact (if any) with
the loreal usually shorter than the contact of the first medial supra-
ocular with the loreal. Two or three suboculars; three, rarely four,
postoculars. Four primary and four secondary temporals, the
uppermost primary in contact with tlie uppermost secondary, the
•owest primary in contact with the two lowest secondaries and with
the penultimate and antepenultimate supralabials. Supralabials
9-11, usually ten; infralabials 6-9. Postmental paired, follow^ed by
three large pairs and one smaller pair of chin shields, of which the
members of the first pair are in contact along the mid-ventral line.
The sublabials extend anteriorly to the first chin shields and second
infralabials, occasionally to the postmental.

Dorsals in fourteen longitudinal rows, frequently only twelve in
specimens from the Distrito Federal and westward; 34-42 (ave.
39.3) transverse rows. Ventrals in twelve longitudinal and 50-58
(ave. 55.1) transverse series. Caudal whorls 91-108 (ave. 100.5).
The median six or eight rows of dorsals strongly keeled, with one or
two lateral rows more weakly keeled.

Tan to brown above, males nearly uniform or with a few small
scattered white spots. Females with dark brown vertical bars on
the sides, sometimes bordered posteriorly with white; a correspond-
ing series of mid-dorsal dark blotches, frequently expanded into bars
more or less confluent with those of the sides. Ventral color yellow-
ish to dark gray, darker laterally than medially. There is no
mottling or prominent spotting, but in a few specimens there is a
tendency for the dark color to form transverse lines between the
scale rows. Infralabials and sublabials usually rather dark, but
chin shields usually light excepting in specimens from Veracruz and
northern Puebla. Females from the western part of the range may
have a color pattern similar to that of the male, with the dark bars
very indistinct or lacking.

These are moderately large lizards, adults averaging about 110
mm. in snout-vent length, the maximum observed being 126 mm.
16—3659

242 The University Science Billf:tin

Tail about 1.35-1.50 times tiie snout- vent length, slightly longer in
the younger specimens than in the older ones.

Remarks: The species imbricata is well defined and readily dis-
tinguishable from both rudicollis and levicollis, though more closely
related to the latter. Probably the most convenient character is the
number of transverse dorsal scale rows, 34-45 in imbricata, below 33
in rudicollis, and above 45 (with a single known exception) in levi-
collis. Geographic races within the species are, however, not so
easily determined; only one such race (/'. ciliaris) is really well de-
fined. In the case of variants found near the ]icriphery of the
known range, especially where the boundary of the known range
probably represents a boundary of an area worked by collectors
rather than a boundary of the true range of the animal, the question
of whether or not truly subspecific populations are represented by
the known specimens cannot be definitely answered at present.
Specific cases of this sort will be mentioned subsequently.

The majority of known specimens of i. imbricata are from the
vicinity of the Distrito Federal, but I do not believe these are entirely
typical of the subspecies. The typical populations inhabit eastern
Mexico and Morelos, southern Pucbla, and northern Oaxaca. These
specimens all have fourteen dorsal scale rows, uniformly light chin
shields, no canthal, and an anterior superciliary which, while not
large, separates the preocular from the first medial supraocular.

Specimens from western Mexico and Michoacan represent a peri-
l^heral population of the ty])e mentioned above. For this popula-
tion Dr. Hobart M. Smith recently (1942) revived the name of
aclspersa Wiegmann, 1834. With his permission, I offer the follow-
ing comments, taken from his unjniblished notes, which he has
kindly allowed me to make use of: "(r. adspersus generally has been
considered a synonym of imbricatus. It appears that Wiegmann
was at the point of describing it when Wagler jniblished the de-
scription of lichenigerus. Wiegmann concluded that his adspersus
was a synonym of lichenigerus, and so treated it in his Herjietologia
Mexicana (1834). However, a figure of the head plates of adsper-
sus was published, under that name, in the latter work. The figurf3
shows that the nasal is broadly in contact with tiie rostral, a condi-
tion which very rarely occurs in imbricnttis. Wiegmann's descrip-
tion of lichenigerus, probably based solely upon his type of ad-
spersus, states that the longitudinal series of dorsals number 12."
Smith had available a single specimen from western Mexico (state) ;
this specimen has twelve dorsal scale rows and the nasal in contact
with the rostral, agreeing in these respects with adspersa, and in

Tihen: Lizard Genus Barisia 243

view of the extremely small probability of finding these two charac-
teristics together on the basis of pure chance, concluded that a west-
ern subspecies exists to which the name adspersa is applicable. I
have since had the opportunity to examine one more specimen from
western Mexico and three from Michoacan, making a total of five
specimens from the presumed range of this questionable form. Of
these, two possessed a full fourteen scale rows; only the one showed
a naso-rostral contact (a condition seen on only one side of one in-
dividual from any other part of the range of inibricata) , although in
another the separation was decidedly narrower than usual ; three of
the five had a canthal split off from the lorcal on at least one side —
another condition rarely found in specimens from the central part
of the range and also exhibited in Wiegmann's figure of adspersa.
In one female specimen the transverse bands are indistinct, in the
other (three are males) these bands are indiscernible, indicating a
possibility of a lesser sexual dimorphism in color jiattern than in the
typical population. The occurrence of all these variants certainly
shows that this western population is atypical, but, on the basis of
the few specimens available, none show a high enough percentage
of occurrence to allow definition of a recognizable subspecies in this
area; it may simply be a case of "peripheral variation." I there-
fore consider it best for the present to continue to consider adspersa
as a synonym of inibricata. By this I do not imply that I do not
believe it ciuitc possible that a recognizable subspecies is repre-
sented, but that present specimens arc insufficient to demonstrate
the existence of such a subspecies, or to provide a diagnosis. If fur-
ther collection shows that this population is demonstrably subspe-
cifically distinct, the name adspersa would probably be correctly
applied to it. It should be mentioned here that a relatively large
number of specimens from the Distrito Federal and vicinity have
only twelve dorsal scale rows, and might in this respect be thought
of as "intergrades."

A series from Cruz Blanco, Veracruz, has the preocular and first
medial supraocular in contact in about 60% of the specimens, and
the chin shields are gray or marked with gray in all adults. Spe-
cimens from Orizaba and Acultzingo to the south and Teziutlan,
Puebla, to the northwest, exhibit these same features to a lesser
degree. Although this geographic variant is well marked, I do not
believe that the proposal of a new subspecific name is warranted,
terming, as would be necessary if such a course were followed, the
Acultzingo-Orizaba and Teziutlan populations as intergrades be-
tween the Cruz Blanco and typical forms.

244 The University Science Bulletin

Specimens examined: Distrito Federal: Near Ajusco (AMNH 15486); Chiribusco (MCZ
28377); near Guadelupe (AMNH 15483); near Mexico (city) (EHT-HMS 10406); near
San Juanico (AMNH 15484-15485); San Juan Teotihuacan (MCZ 6345, 4 spec).

Mexico: Desierto de los Leones (EHT-HMS 10398-10401); Lake Lerma (EHT-HMS
19144-19145); Llano Grande (EHT-HMS 23838-23840, USNM 113213); Nevado de Toluca
(EHT-HALS 22218-22219, 23808-23809, 23830-23837; USNM 113219); San Martin (EHT
10396); Villa Victoria (USNM 113215); Zempoala (USNM 113209-113212).

Mexico or Morelos: Lake Zempoala (EHT 10402-10404; 10426, 19133-19136); Monte
Rio Frio, 45 km. ESE of Mexico (city) (TAM 711, 738-743); 55 km. SE of Mexico (city)
(TAM 911-918); 45 km. SW of Mexico (city) (TAM 919-920).

Michoacdn: Mount Tancitaro (USNM 47787, UMMZ 94411); 10 km. Northwest of
Zacapu (UMMZ 94412).

Morelos: Tepoztlan (EHT-HMS 28870); near Tres Marias (EHT-HMS 10422).

Puebia: Cacoloapam (EHT-HMS 10397); Teziutlan (KHT-HMS 23841-23840); Popo-
catepetl (EHT-HMS 19137, 23807); Rio Otlati, 15 km. NW of San Martin (TAM 901).

Veracruz: Acultzingo (USNM 113204-113208, EHT-HMS 28871-28878); Cruz Blanco
(EHT-HMS 10405, 10407-10417, 10423-10425, 10427); Orizaba (EHT-HMS 19138-19139,
USNM 7087, 2 spec, 19262, 47774); 4 km. S of Jalacingo (Univ. of Kansas, uncatalogued).

Hidalgo: EI Chico National Park (EHT-HMS 19141); San Miguel (MCZ 11434);
Tianguistengo (EHT 23848); Zacualtipan (EHT-HMS 19142, 23847).

Oaxaca: Reyes (USNM 47392); Zempoaltepec (?) (47189-47191).

Indefinite or extremely doubtful localities: USNM 3088, 7036, 8317, 26165, 32166;
AMNH 19304, 24760-24763.

A total of 128 specimens. The form is also reported from Palamba, Michoacan; Cuer-
nevaca, Morelos; and Nevado de Colima, Jalisco, the last not certainly referable to this form.

Barisia imbricata ciliaris (Smith)

Gerrhonotus levicolti-s ciliaris Smith, 1942, Proc. U. S. N. M., 92, no. 3153: 365.

Type and type locality: USNM 47496; from Sierra Guadelupe,
Coahuila.

Range: Hidalgo and Guanajuato northward to Chihuahua and
Coahuila.

Diagnosis : A Barisia of the imhricata group with twelve rows of
ventrals; one loreal, usually no canthal; nasal separated from the
rostral; about 39-45 transverse and usually sixteen longitudinal
rows of dorsals; dorsal color a nearly uniform light brown above in
both sexes, often spotted with white; venter light; first superciliary
moderate or large, separating the preocular and first medial supra-
ocular; lowest primary temporal in contact with the penultimate
and antepenultimate supralabials.

Description: Nasal separated from the rostral b}' the inter-
nasals. One loreal; normally no canthals; postnasal in rather broad
contact with the supranasal. Three superciliries, the anterior nor-
mally separating the preocular from the first medial supraocular;
first superciliary moderately large, its contact with the loreal usu-
ally about equal to the contact of the first medial supraocular with
the loreal. Two suboculars; three postoculars. Four primaiy and
four secondaiy temporals, the ujipermost primary in contact with
the ui)permost secondary, the lowest ]irimary in contact with the

Tihen: Lizard Genus Barisia 245

two lowest secondaries and with the penultimate and antepenulti-
mate supralabials. Supralabials 9-11; infralabials 8-10. Postmen-
tal paired, followed by three large pairs and one smaller pair of chin
shields, of which the members of the first pair, occasionally also the
second, are in contact along the mid-ventral line. The sublabials
usually extend anteriorly to the first chin shields and second infra-
labials, frequently to the postmental, less commonly to the first in-
fralabial.

Dorsals usually in sixteen longitudinal and 39-45 (ave. 41.3)
transverse rows; Smith records a single specimen with 49 transverse
rows, but I have seen none with more than 45. Ventrals in twelve,
occasionally fourteen, longitudinal and 53-61 (ave. 57.8) transverse
rows. Caudal whorls 87-102. The median six or eight rows of dor-
sals are keeled.

A nearly uniform light brown above, often with a reddish or
yellowish cast; small scattered white spots often present. Venter
uniformly light except laterally, where it may be slightly darkened.

These are large lizards, adults averaging about 120 mm. in snout-
vent length; the maximum observed is 145 mm. Tail about 1.30-
1.60 times the snout-vent length.

Remarks: This form is distinguished from i. imhricata by the
larger number of dorsal scale rows, both transverse and longitudinal,
by the color pattern (particularly that of the f-emale), and to a
lesser extent by the larger first superciliary. This is also a definitely
larger race.

A female specimen from Atotonilco Grande, Hidalgo resembles
i. imhricata in scalation, but has the uniform color of ciliaris. Other
southern Hidalgo specimens resemble i. iinhricatus in all respects,
while a specimen from northern Hidalgo is typical of ciliaris. A
subspecific relationship between the two forms, with intergradation
occurring in central Hidalgo, appears probable.

A female specimen (LSU R28) has an unpaired postmental; four
of ten embryos contained in the uteri have this same rare condition.

Specimens examined: Durango: Inde (USNM 46843).

Guanajuato: San Felipe (EHT-HMS 10418-10421).

Hidalgo: Durango (EHT-HMS 2380.3). ciliaris x imhricata, Atotonilco Grande (EHT-
HMS 19140).

Ntievo Leon: Pablillo, Galeana (EHT-HMS 19131).

San Luis Potosi: No further data (MCZ 4547, 8339); near Jesus Maria (USNM 47207-
47210); between Llano de Coneja and Llano de Garzas, 7,000 ft. (R28 Louisiana State Univ.).

Sinaloa: Esquinapa (AMNH 585).

A total of sixteen specimens. Other locality records are: Sierra Guadelupe, Coahuila;
Coyotes, Durango; Sierra de Santa R-osa, Guanajuato; Sierra Madre, Zacatecas.

246 The University Science Bulletin

Barisia imhricata planifrons (Bocourt)

Gerrhonotus (Barissia) planifrons Bocourt, 1879, Miss. Sci. Mex., Rept., livr. 6: 301.
Gerrhonotus planifrons, Giinther, 1885, Biol. Centr. Am., Rept. : 38.
Barissia planifrons, Cope, 1887, Proe. Amer. Philos. Soc, 17 : 97.

Type and type locality: A presumably young specimen in the
Paris Museum; the type locality is given only as "Oaxaca."

Range: Uncertain, but ])robably southern Oaxaca; typical /. im-
bricata is known from Reyes, and two specimens from Cerro San
Felipe are thought to be imhricata X planifrons intergrades.

Diagnosis. A Barisia of the imhricata group with twelve rows of
ventrals; one loreal and usually no canthal; nasal separated from
the rostral; about 35-39 transverse and sixteen longitudinal rows of
dorsals; coloration as in i. imhricata; first superciliary relatively
large, separating the preocular and first medial supraocular; lowest
primary temporal in contact with only the antepenultimate supra-
labial.

Description: The following description applies to the type speci-
men, as described and figured by Bocourt.

Nasal separated from the rostral by the anterior internasals. One
loreal; no canthal; postnasal slightly separated from the supranasal
by a naso-loreal contact. Three superciliaries, the anterior sei^a-
rating the preocular and first medial supraocular; it is of large size,
its contact with the loreal greater than the contact of the first
medial supraocular with the loreal. Two suboculars; three post-
oculars. Four ]irimary and four secondary temporals, the upper-
most primary in contact with the uppermost secondary, the lowest
primary in contact with the two lowest secondaries and with the
antepenultimate, but not the jienultimate, supralabial. Supralabials
ten; infralabials eight or nine. Postmental paired, followed by three
large pairs and one smaller pair of chin shields, of which the mem-
bers of the first pair are in contact along the mid-ventral line.

Dorsals in sixteen longitudinal and about thirty-five transverse
rows; ventrals in twelve longitudinal rows. The median eight rows
of dorsals are strongly keeled.

Color pattern apparently similar to that of B. i. imhricata.

Remarks: Two specimens from Cerro San Felipe, Oaxaca, agree
with the above description except as follows: One specimen has a
canthal split from the dorsal portion of the loreal on one side ; supra-
nasal and postnasal in contact in both ; in one the lowest primary
temporal is in contact with the ])enultimate supralabial; dorsals in
sixteen longitudinal rows in one, fourteen to fifteen in the other; ven-
trals in 53 and 54 transverse rows.

Tihen: Lizard Genus Barisia 247

Though these specimens are not entirely typical of planifrons as
described above, they do resemble the type specimen of that form
in certain features. I have therefore tentatively assigned them to
the category of inibricata X planifrons, assuming that a population
to which the name planifrons is applicable actually exists some-
where south of Cerro San Felipe.

In many respects this form resembles ciliaris more closely than it
does typical imhricata, but differs in the lower number of transverse
dorsal scale rows and in the fact that the lowest primary temporal
is not in contact with the penultimate supralabial, probably also in
color pattern.

Specimens examined: Oaxaca: Cerro San Felipe, planifrons x imhricata intergrades (EHT-
HMS 19132, 19143). No other specimens known witli definite locality data.

Barisia levicollis Stejneger

Barissia levicollis Stejneger, 1890, Proc. U. S. Nat. Mus., 13, no. 809: 184.

Gerrhonotus imbricatus levicollis, Dunn, 1930, Proc. Acad. Nat. Sci. Philad., 88: 475.

Gerrlwnotus levicollis levicollis. Smith, 1942, Proc. U. S. Nat. Mns.. 92, no. 31.53: 308.

Type and type locality: USNM 9362; said to be from the ":\Iexi-
can boundary/' probably from central Chihuahua.

Range: Known only from central and southern Chihuahua.

Diagnosis: A Barisia of the inibricata group with twelve rows of
ventrals; one loreal and no canthal; nasal separated from the ros-
tral; forty-six or more transverse and sixteen longitudinal rows of
dorsals; uniform light brown above, with or without small scattered
white spots; venter light, often with black spots laterally; a single
(middle) superciliary, the preocular in contact with the first medial
supraocular; lowest primary temporal usually in contact with only
the antepenultimate supralabial.

Description: Nasals separated from the rostral by the anterior
internasals. One loreal, often separated from the prefrontal by a
contact of the first medial supraocular with the posterior internasal ;
no canthals. Postnasal in rather broad contact with the supranasal,
usually also with the posterior internasal. Only a single super-
ciliary present, corresponding to the most posterior element of this
series in imhricata; the preocular therefore in contact with the first
medial supraocular, and usually also with the first lateral supra-
ocular. Two suboculars; two postoculars, the upper quite large, the
lower very small. Four primaiy and four secondary temporals, the
uppermost primary in contact with the uppermost secondary, the
lowest primary in contact with the two lowest secondaries and with
the antepenultimate but not the penultimate supralabial. Supra-
labials 9-11; infralabials 6-9. Postmental parired, followed by three

248 The University Science Bulletin

large pairs and one smaller pair of eliin shiekls, of which the mem-
bers of the first pair are in contact along the mid-ventral line. The
sublabials extend anteriorly to the first chin shields, occasionally the
postmental, and the second infralabials.

Dorsals in sixteen longitudinal and 45-51 (ave. 47.8) transverse
series; ventrals in twelve longitudinal and 57-64 (ave. 61.0) trans-
verse rows. Caudal whorls 99 and 102 in two specimens with un-
damaged tails. The median six or eight rows of dorsals are keeled,
but not so strongly as in imhricata.

Both sexes a uniform tan above, occasionally with small scattered
white spots. Venter nearly uniform yellowish or liglit gray, usually
with small blackish spots laterally.

This is the largest fomi of the genus, adults averaging about 125
mm. in snout-vent length ; the observed maximum 157 mm. Tail
about 1.4 times the snout-vent length.

Remarks: Des})ite the fact that in certain respects ciliaris is in-
termediate between /. imhricata and levicollis, the latter is possessed
of so many unique characters that I have chosen, pending further
information, to consider it specifically distinct. The range of levi-
collis within Chihuahua is not known, since most of the si>ecimens
bear incomplete locality data, but it certainly extends as far south
as Batopilas. All of the specimens I have seen are typical levicollis,
with no tendency towards ciliaris. A single specimen of ciliaris with
the only locality data being "Chihuahua" is completely typical of
that form. Thus while there is no positive evidence of any overlap
in the ranges of these two forms, there is at least some indication
that such an overlap may occur. The Chihuahua-Durango border
region would appear to be the critical area for the solution of this
problem.

Speciynens examined: Chihuahua: No furtlier data (AMNH 1945, USNM 2C603) ; "north
Chihuahua" (AMNH 592-594); Colonia Garcia (MCZ 6977); Meadow Valley (USNM 26602;
2fir,i2); 65 miles east of Batopilas (USNM 47413).

A total of nine specimens. Also reported from Samachique, Chihuahua.

Tihen: Lizard Genus BARifiiA 249

GENERAL REMARKS

I have previously (American Midland Naturalist) pointed out
that the Barisia-Gerrhonotus-Elgcma ancestral stock probably in-
habited the northern Mexican plateau region in the lower Oligocene,
and postulated possession by this stock of the following character-
istics: Skull similar to that of Elgaria; dorsal osteoderms as in
Barisia; cephalic scutellation much as in Gerrhonotus I. infer-
nolis; body scales moderately numerous, extensively carinated, much
as in Barisia moreleti; general color pattern similar to that of Bar-
isia moreleti and Elgaria c. coendea, i. e., a longitudinal light band
dorsally and black white-tipped scales laterally, irregularly ar-
ranged; belly rather strongly mottled; postero-ventral surfaces of
forearms and shanks nearly agranular; tail moderate in length.

One section of this stock became mochfied at the center of disper-
sal; the pter\'goid teeth were reduced, there was some reduction of
the loreo-canthal group of head plates, and the posteroventral sur-
faces of the forearms and shanks became more granular. These and
other minor modifications produced a ''Barisia prototype" stock in
this central area. Continued modification in and southward migra-
tion from this area produced the present arrangement of forms
within the genus Barisia, as discussed in the subsequent paragraphs.
There was no northward migration of Barisia from this region, but
the establishment of a proto-Elgaria stock in the Chihuahua-Sonora
region is probably an integral part of the same general outward
movement as the southward migration of the early Barisia.

As indicated, the Barisia prototype was similar in many respects
to B. m. moreleti, but with certain important exceptions. The
pterygoid teeth, though probably in the process of reduction, had
not been completely lost; the internasal, frontonasal, and prefrontal
elements were all moderately developed; a postrostral was present,
at least in a certain proportion of this stock.

Certain specializations appeared in the population near the center
of dispersal, while forms retaining the more primitive characters
migrated southward. The first migrant group crossed the Isthmus
of Tehuantepec, becoming established south of that barrier, but the
movement did not proceed beyond northern Central America at that
time; this may have been due to an inability to overcome the barrier
constituted by the southern Nicaragua lowlands. That area was
submerged in the early and middle Miocene, but complete submer-
gence was probably not necessary to form an effective barrier.

The single postmental, characteristic of the moreleti group, was
probably developed during the period of southward movement, since

250 The University Science Bulletin

it is found in viridiflava. This species is thought to have originated
from a portion of the group which survived north of the Isthmus,
rather than one which re-entered the area from the south. The only
speciaHzed group characters in which it agrees with the southern
members are the complete lack of pterygoid teeth and the possession
of an unpaired postmental. The usual absence of a frontonasal and
the frequent occurrence of a postrostral indicate that separation
from the remainder of the group took place before the modifications
affecting these elements in the rest of the group occurred. INIore-
over, a large number of modifications are found in viridiflava but
not in the more southern members of the group; this might also be
taken as evidence for a long period of isolation of this species.

The Guatemala-Nicaragua section developed an enlarged fronto-
nasal and lost the postrostral; these modifications are common to all
races of moreleti (and also to monticola) . In different jiarts of the
range the minor modifications appeared which differentiate the
existing subspecies of moreleti, as have been mentioned in the text.

The entrance of members of this group into southern Central
America was subsequent to the establishment of the species moreleti
in the north, rather than an integral part of the original southward
movement. All of the specializations common to all races of more-
leti are also to be found here, in addition to those others which have
appeared subsequent to separation from the northern group. These
latter include particularly a modification of the color pattern and a
reduction of the number of longitudinal dorsal scale rows. The
number of transverse dorsal rows is also somewhat reduced, and the
supranasal has been lost in a large proportion of the population.

The second group moving southward from the dispersal center
differed in a number of respects from the first. Here again the
pterygoid teeth were completely lost, but the postmental in this case
remained paired. The color pattern was slightly modified, the dark,
usually white-tipped, scales of the sides being arranged into definite,
more or less regular, vertical bars, which were sometimes continued
across the back. A fronto-premaxillary contact developed in the
skull ; the number of longitudinal dorsal scale rows was reduced to
sixteen, and the general size increased slightly. All of these modifi-
cations, with the exception of the complete loss of the pterygoid
teeth, must have occurred in the original center of dispersal, rather
than during the course of migration, since they are shared by this
group and the succeeding one which originated from that center.

This second migrant group, the prototype of the gadovii group,
pushed southward to the Isthmus of Tehuantepec but failed to cross

Tihen: Lizard Genus Barisia 251

that barrier. It has since been obliterated everywhere except for a
small area in southern Guerrero and northern Oaxaca, and another
limited area in Veracruz. The Guerrero form, B. g. gadovii, is
thought to resemble very closely the original migrant population,
although there is a loss of certain elements, such as the postrostral,
from a large percentage of the population. The Oaxacan g. levigata
shows only minor modifications of coloration and scalation. The
Veracruz group reduced the number of loreo-canthal elements, ex-
panded the supranasals, and in the case of antauges presumably lost
the frontonasal.

The population remaining at the center of dispersal, in contrast to
these two major migrant groups,, retained pterygoid teeth in a ves-
tigial condition. Numerous modifications in scalation appeared, in
addition to those shared with the gadovii group. The posterior
superciliaries, frontonasal, upper postnasal and postrostral were all
lost. The granular area of the lateral fold was decidedly enlarged.
The entire color pattern was greatly lightened. The loreo-canthal
group was reduced to probably two loreals and a single (posterior)
canthal on each side. There was also a general increase in size.

A portion of this population became somehow isolated at a rela-
tively early date, yet without being geographically far removed
from the center of dispersal. This particular form, rudicollis, be-
came highly specialized in several respects, e. g., the great reduction
in number of transverse dorsal scale rows, but the retention of the
anterior loreal and of four superciliaries indicate that it is an earlier
derivative of the imbricata stock than are other existing members
of the group. The early distribution of this form may liave been
somewhat different than at present, but evidence has been presented
that its early isolation may have been ecological rather than geo-
graphical.

In the central population the number of superciliaries was reduced
to three and the anterior loreal lost. Some such form as this spread
over most of Mexico south to, or nearly to, the Isthmus of Tehuan-
tepec. In the central to southern part of that range the anterior
superciliary was somewhat reduced and the niunber of dorsal scale
rows reduced to fourteen. These two features spread throughout
most of the region south of 21 degrees latitude, except for a small
area in Oaxaca where planifrons presumably occurs. In the western
part of that area the number of dorsal scale rows has been further
leduced to twelve in a sizable percentage of the population, and the
anterior canthal perhaps retained in a relatively large proportion.
Throughout the remainder of the range, including the area north of
21 degrees, the canthal was lost in the majority of the population.

252 The University Science Bulletin

Roughly north of 21 degrees these changes did not occur, except
that in cilinris the number of transverse rows of dorsals may have
decreased slightly. The northern forms are definitely larger than
the southern, probably larger than the early widespread population.
In addition, the dark transverse bars have been lost in both sexes.

In Chihuahua the high number of transverse dorsal rows was re-
tained. As in all the northern group, the dark bars were lost in both
sexes and the size presumably increased. Other modifications have
occurred here, the most obvious being the reduction of the super-
ciliary series to a single element. Others include the usual contact
of the posterior internasals with the postnasals, and often also with
the first medial supraoculars.

Although all forms of this genus are found only at relatively high
altitudes, it appears that members of the moreleti group regularly
occupy somewhat higher altitudes, on the average, than do members
of either of the other groups. Certain characteristics of that group
should be pointed out as probable correlatives of that fact. Firstly,
the individuals in this group are decidedly smaller than are repre-
sentatives of the other two groups. Secondly, this group is more
heavily and darkly pigmented than the other groups, the difference
from the gadovii group, however, being much less pronounced than
that from the imbricata group. This dark coloration and small size
are frequent characteristics of poikilothermous forms found at
high altitudes (or northern latitudes), the presumption being that
these features allow more efficient utilization of radiant heat from
the sun, the dark pigment increasing the absorptive capacity of
the surface and the small size producing a greater surface area
relative to the mass of tlie body. Thirdly, it seems probable
that the moreleti group is ovoviviparous, the other groups may
or may not be. I know of no direct observations on this j^oint
for any of the forms concerned, but eggs have been observed
in the uterus of female specimens of the moreleti group with em-
bryos up to 26 mm. in length; it is therefore obvious that consider-
able development takes place while the egg remains in the uterus,
and it is probable that the young are born alive. In the other two
groups, well-developed embryos were found only in B. i. ciliaris;
fully matured eggs were frequently found in other forms, but no
recognizable embryos were ever observed. In both gadovii and i.
imbricata a series of specimens taken at the same time (approxi-
mately) contained females with mature eggs and one or more fe-
males in which the uterus was empty and had a deflated appearance,
indicating that oviposition had recently occurred. This would in-
dicate an oviparous habit for these forms.

Tihen: Lizard Genus Barisia 253

LITERATURE CITED

Agassiz, Louis. 1846. Nomenclatoris zoologici index universalis. Soloduri,

1846. pp. i-viii, 1-393.
Baird, Spexcer F. 1859. Description of new genera and species of North

American lizards in the museum of the Smithsonian Institution. Proc.

Acad. Nat. Sci. Philadelphia, for 1858, vol. 10: pp. 253-256.
BocouRT, M. F. 1871. Description de quelques Gerrhonotes nouveaux pro-

venant de Mexique et de I'Amerique Centrale. Nouv. Arch. Mus., 7 (Bull) :

1871, pp. 101-108.
1873. Notes erpetologiques. Ann. Sci. Nat., ser. 5, vol. 17, no. 2:

not paged.

1878-1879. Livraison 5-6 (pp. 281-440. pis. 20, D-G, 21, A-C, 22, 22,

A-D) in Dumeril, Bocourt and Mocquard, fitudes sur les reptiles. Miss.
Sci. Mex., Zool., 3 (sec. 1), pp. i-xiv, 1-1012.
BouLENGER, George Albert. 1913. Description of new lizards in the collec-
tion of the British Museum. Ann. Mag. Nat. Hist., ser. 8, vol. 12, 1913,
pp. 563-566.

Cope, Edward D. 1866. Fourth contribution to the herpetologv of tropical
America. Proc. Acad. Nat. Sci. Philadelphia, vol. 18 (2). 1866, pp. 123-132.

1875. Check-list of North American batrachia and reptilia; with a

svstematic list of the higher groups, and an essay on geographical distri-
bution. Bull. U. S. Nat. Mus., no. 1, 1875, pp. 1-104.

1877. Tenth contribution to the herpetology of tropical America.

Proc. Amer. Philos. Soc, vol. 17, 1877, pp. 85-98.

1887. Catalogue of batrachians and reptiles of Central America and

Mexico. Bull. U. S. Nat. Mus., no. 32, 1887, pp. 1-98.

1900. The crocodilians, lizards and snakes of North America. Report

U. S. Nat. Mus. for 1898 (1900), pp. 153-1270.

Dunn, Emmet Reid. 1936. The amphibians and reptiles of the Mexican
Expedition of 1934. Proc. Acad. Nat. Sci. Philadelphia, vol. 88, 1936, pp.
471-477.

and John T. Emlen, Jr. 1932. Reptiles and amphibians from Hon-
duras. Proc. Acad. Nat. Sci. Philadelphia, vol. 84. 1932, pp. 21-32.

FiTziNGER, L. I. 1843. Systema reptilium. pp. 1-106.

Gr.\y, John Edward. 1838. Catalogue of the slender-tongued saurians, with

descriptions of manv new genera and species (part 2). Ann. Mag. Nat.

Hist., ser. 1, vol. 1, 1838, pp. 388-394.

— ■ 1845. Catalogue of the specimens of lizards in the collection of the

British Museum. London, pp. i-xxviii, 1-289.

GtTNTHER, Albert C. L. G. 1885-1902. Biologia Centrali-Americana. Rep-
tilia and Batrachia. pp. 1-326, pis. 1-76.

H.ARTWEG, Norman and Joe A. Tihen. 1946. Lizards of the genus Ger-

rhonotus from Chiapas. Occ. Pap. Univ. Mich. Mus. Zool., no. 497, 1946,

pp. 1-16.
Martin del Campo, Rafael, 1939. Contribucion al conocimiento de los Ger-

rhonoti Mexicanos, con la presentacion de una nueva forma. Anal. Inst.

Biol., vol. 10, pt. (3/4), 1939, pp. 353-361.

Peters, W. 1877. Eine Mittheilung iiber neue Arten der Sauriergattung
Gerrhonotus. Monatsber. Konig. Prcus. Akad. Wiss. Berlin for 1876 (1877),
pp. 298-300.

Schmidt, Iv.\rl P. 1928. Reptiles collected in Salvador for the California
Institute of Technology. Field Mus. Nat. Hist. Zool. Ser., vol. 12, no. 16:
pp. 193-201.

254 The University Science Bulletin

Smith. Hobart M. 1942. Mexican heipctological mi.«cellanv. Proc. U. S.

Nat. Mus., vol. 92, no. 3153, 1942. pp. 349-395, pi. 37.
Stbjneger, Lex)nhard. 1890. On the North American lizards of the genus

Barissia of Gray. Proc. U. S. Nat. Mus., vol. 13, no. 809, 1890, pp. 183-185.

1907. A new gerrhonotine lizard from Costa Rica. Proc. U. S. Nat.

Mus., vol. 32, no. 1542, 1907, pp. 505-506.

Stu.\rt, L. C. 1943. Comments on the herpetofauna of the Sierra de los
Cuchumatanes of Guatemala. Occ. Pap. Univ. Mich. Mus. Zool., no. 471,
1943, pp. 1-28, pi. 1.

SuMiCHRAST, F. 1882. Enumeracion de las especies de reptiles observados
en la parte meridional de la Republica Mexicana. La Naturaleza, vol. 6,
1882, pp. 31-45.

TiHEN, J. A. (in press). The genera of gerrhonotine lizards. Amer. Mid.

Nat. 1949.
W.\GLER, J. 1830. Descriptiones et icones amphibiorum. Fasc. II.

WiEGMANN, A. F. 1828. Bevtrage zur Amphibienkunde. Isis. no. 3, 1828,
pp. 364-383.

1834. Herpetologia Mexicana. Pars prima, saurorum species. Berlin.

pp. 1-51.

Tihen: Lizard Genus Barisia

255

Plate I. Barisia rudicollis. Specimen in Institute de Biologia de
Mexico. Hacienda de la Gavia, Mexico (state) Mexico. Actual snout-
co-vent length, 105 mm.

■mistmsmrmmmmi:^,^,..,.^

-•O {^^

-s^i^fj^ jfc»»»*

Plate II. Figures 1 and 2. Ventral and dorsal views of Barisia mon-
iicnln. Museum of Comparative Zoology. Harvard College no. 15467;
Volcan Poas, Costa Rica. Actual snout-to-vent length, 75 mm. Figures
3, 4, and 5, BariHia gndovii hviijuta. Holoty{)e U. S. National Museum
no. 47212; "Valley of Oaxaea" Mexico.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. I] April 20, 1949 [No. 4

Costa Rican Frogs of the Genera Centrolene and

Centrolenella

BY

Edward H. Taylor

Abstract. — Four species belonging to the genera Centrolene and Centrole-
nella have been recognized in the Fauna of Costa Rica. These are Centrolene
■prosohlepon (Boettger), Centrolenella pulveratum (Peters), Centrolenella
valerioi (Dunn), Centrolenella fleischmanni Boettger. Four forms are de-
scribed as new in the genus Centrolenella. These are C. spinosa, C. granulosa,
C. albomaculata and C. colymbiphyllum, all from Costa Rica.

Twenty-five specimens of small, green, hylid frogs taken in Costa^
Rica in the summer of 1947, resolve themselves into a group of
species, one of which is referable to the genus Centrolene, since a
humeral protuberance is present, and the others to the genus Cen-
trolenella in which the protuberance or hook is absent. One of the
species possesses a rudiment of a pollex with a sharp spine usually
penetrating the free tip. While I am inclined to regard this develop-
ment as worthy of generic designation, I am for the present referring
the species to Centrolenella pending further research in the matter.

Three species have already been described from Costa Rica.
These are Hyla prosoblepon Boettger, in 1892, from Plantage Cairo
(La Junta) near Limon, Atlantic side of Costa Rica; Carl Fleisch-
mann collector; Hylella fleischmanni Boettger, in 1893, from San
Jose, Costa Rica, Carl Fleischmann collector; and Centrolene valerioi
Dunn, in 1931, from La Palma, Costa Rica, Manuel Valerio and E.
R. Dunn collectors. A fourth species has been reported in Costa
Rica. This is Hyla pulverata Peters, described in 1873, from Chiri-
qui, Panama, H. Ribbe, collector. The present designations of the
four species occurring in Costa Rica are: Centrolene proseblepon

17—3659

(257)

258 The University Science Bulletin

(Boettger), Centrolenella pulveratum^ (Peters), C entrolenella val-
erioi (Dunn) and Centrolenella fieischmanni (Boettger).

A free translation of the original description of Hijla pulverata
Peters follows:

Vomerine teeth in two groups between the choanae which are much larger
than the small openings of the Eustachian tubes. Tongue scarcely emarginate
behind. Snout, not longer than the eye, with a rounded canthus rostralis.
Tympanum round, very small, and covei'ed over with pigmented skin. Back
smooth, the belly and vmderside of thigh granular; first and second fingers
apparently equally long, much shorter than the fourth; first finger entirely
free; the web between the second and third fingers reaches almost to the
terminal pads of the second and somewhat over the base of the first joint of
the second; between the third and fourth from the terminal disk of the fourth
to the middle of the penultimate phalanx of the third. The toes are bound
with a complete web which leaves only the two outer joints of the fourth toe
free.

Colorless, with scattered small white punctations which are numerous on
the side of the head. Under a lens the surface is seen to be punctate with
dark flecks.

Total length in mm., 24; head, 9; head width, 8; arm, 15; hand, with third
finger, 8; leg, 45; foot and fourth toe, 20.

One specimen from Chiriqui, from H. Ribbe. This species in its form and
tooth arrangement is nearest Hlyla] rodopepla, while its color recalls H.
punctata and albomarginata. The very well-developed web between the fingers
and the very small tympanum without a skinfold over it serves to separate
them easily.

I have not been able to recognize this species in any of my Costa
Rican material. Among the specimens at hand are several species,
vv^hich do not conform to either structures or color of the described
forms and I conclude that they represent undescribed species, and
are here described.

Centrolene prosoblepon (Boettger)

Hyla prosoblepon Boettger, Katalog der Batrachier — Sammlung in Museum der Sencken-
bergischen Naturforschenden Gesellschaft in Frankfurt am Main, Aug., 1892, p. 45. (Type
locality, Plantage Cairo [La Junta], near Limon, Atlantic side of Costa Rica): Giinther,
Biologia Centrali -Americana, Reptilia and Batrachia, Sept., 1901, p. 280.

tHyla puncticrus Boulenger, Ann. and Mag. Nat. Hist., ser. 6, vol. 18, 1896, p. 341.

Centrolene prosoblepon Noble, Proc. Biol. Soc. Washington, vol. 37, 1925, pp. 66-71.

A series of eight specimens from Isla Bonita (The American Cin-
chona Plantation), chsplay varying shades of lavender, and different
degrees of spotting.

* There is some doubt as to whether there is a humeral hook present in pulveratum.
Peters makes no mention of this organ. This of course may be accounted for if the type is
a female — a strong probability since the vocal sac is not mentioned either.

Dunn, 1931b, who believes he has recognized this form in specimens from El Valle de
Anton and in Co.sta Rica, states that "a rudimentary humeral hook, not projecting" is
present; and later Dunn (1933) in listing characters of the form, states "male and female
without humeral hook." I am presuming that the last statement is correct, and am referring
the species to Centrolenella.

Taylor: Genera Centrolene and Centrolenella 259

Two females, the largest specimens, have the darkest shade of
lavender. The exposed surfaces of the back and limbs have small,
rounded, purplish or purplish-brown spots. The males, which on
the whole are of a lighter shade of lavender, have the spotting similar
to that in the females in two specimens, while in four the spots are
fewer in number and confined to the rump and dorsal surfaces of the
limbs.

The largest male, with a very well-developed humeral process, has
no trace of vomerine teeth but they are present in all other speci-
mens. The two females show no trace of the humeral process.

Noble and Dunn both report that the humeral process is present
in certain females of a species they identify as Centrolene prosoble-
pon. One wonders whether there is an unrecognized species in Cen-
tral America in which this is a constant character in females or
whether the so-called females with the humeral process represent
some type of intersex.

The measurements of the type, as recorded in the type description,
is appended to the following table of measurements. It may be
noted that the measurement of the arm of the type is considerably
smaller than that of specimens of smaller snout-to-vent measure-
ment in this series; the leg is slightly smaller, the head length a
little shorter, and the head width a little greater.

The type is from near sea level while the specimens listed here
come from an elevation of about 5,500 ft. on Volcan Poas. Whether
or not direct comparison with the sea level population would reveal
other significant differences cannot be stated.

Centrolene prosoblepon Boettger

Total
length

Head
length

Head
width

Arm

Hand

and
fingers

Leg

Foot
and
toe

Tibia

Femur

23806

9

27.5

9

9.5

18

8.5

46

20

15

15

793*

5

26.5

9

9.8

16

10

46.2

20

15.2

15

2922

$

26.5

8.4

9.2

18

8

46

19

14.6

13

23804

5.

26

8.3

9

17

8

46

19.4

14.6

13

2926

$

25

8.3

9

17

8

46

19

14.4

14

23803

$

25

8.2

9

16

8

45

19

14.2

13.8

23805

S

■ 24

8

8.5

17.

5

7.9

43

19

14

12.8

794

$

23.4

7.9

8.5

17

7.8

43

19

14

13

Type

24.5

7.5

9

14.

5

42

*Nos. 793, 794,

2922, 2926 are

field nur

nbers

of

R. C. Tf

ivlor.

Centrolenella spinosa sp. nov.

Type: University of Kansas Museum of Natural History no.
23809 ^ . Collected at Los Diamantes, one mile south of Guapiles,
Costa Rica, Sept. -7, 1947, by Richard C. Taylor and Edward H.
Taylor.

260 The University Science Bulletin

Paratypes: Nos. KU 23810, KU 23811; RCT 2467-2469 (all
males). Same locality and collectors. Sept. 7-8, 1947.

Diagnosis: A small species (about 21 mm.) ; snout high, the
front ed^e nearly vertical; nostril near front edge of snout. Vom-
erine teeth very weak, usually present; tympanum present; skin
minutely rugose but not granular on back and sides; a well-defined
pollex rudiment bearing a sharp spine which may or may not be
exposed; lavender without any spotting; eyelids purple; iris silver
with purple reticulation; inner toe narrowed at tip.

Description of the type: Head broader than the body; eyes prom-
inent, directed somewhat forward; canthus rostralis apparent, very
much rounded; areas about nostrils much swollen with a distinct
medial depression between them; loreal region with a long slope
to lip, the medial area behind nostril slightly concave; snout in
front of nostrils nearly vertical, the nostrils much nearer median
point of upper lip than eye; interorbital width greater than the
length of snout, one and three fourths times width of an eyelid;
tympanum present, its greatest diameter about one and three
fourths times in its distance from eye, almost three times in longi-
tudinal diameter of eye; a rather marked constriction of head be-
hind tympanum, with a somewhat swollen area above arm inser-
tion; supratympanic fold obsolete; tongue rather rounded with a
scarcely discernible notch behind, free behind for about one fifth
its length; openings to vocal sacs are elongate curved slits be-
ginning along sides of tongue and extending some distance behind;
choanae large, round, the diameter of one contained in distance
between them three times; external vocal sac forming ample folds
on chin.

Upper arm very slender, the foreann greatly thickened with an
indistinct flattened skin fold on under surface; only a faint trace
of web between the first and second fingers; a small but distin-
tinguishable web between the second and third with a moderate
fringe reaching terminal pad on outside of second finger; third and
fourth fingers about half webbed, the web not reaching the level of
the outer subarticular tubercles; finger widened at tips, and more
or less tmncate ; first finger much longer than second with a distinct
rudiment of pollex, free at tip, with a sharp, strong spine (pos-
sibly absent in female; none seen) ; pollex surrounded by the elon-
gate palmar tubercle; a stongly-defined rounded median palmar
tubercle; subarticular tubercles well defined and numerous; super-
numerary tubercles present; legs long, the tibiotarsal articulation

Taylor: Genera Centrolene and Centrolenella 261

reaching slightly beyond snout tip when limb is adpressed; the
heels touch but do not overlap ; foot a little more than three fourths
webbed, the tei-minal pads smaller than those on fingers, the outer
somewhat truncate, that on inner toe somewhat pointed, scarcely
or not as wide as toe; moderate inner metatarsal tubercle elevated
but little; apparently no trace of an outer tubercle; subarticular
tubercles small, supernumerary tubercles not or scarcely discern-
ible; inner tarsal fold very weak, scarcely discernible; anal opening
high on rump covered by a free flap, the transverse free edge much
longer than width of flap.

Skin on back and sides seen under a lens, minutely rugose, but
definitely not granular; belly and under side of thigh strongly
granular, the granules of the latter much the larger.

Color: In life, yellow green except concealed surfaces of arm,
leg, and under side of body which are flesh white, and transparent,
without markings; a large light cream spot in front of arm inser-
tion. In preservative (formalin, then alcohol) the dorsum nearly
uniform lavender, the eyelids deep purple; limbs lighter but with
a fine peppering of lavender chromatophores on the exposed surface,
even on most of upper surface of the thigh ; below and on concealed
surfaces pure white without pigment; iris of eye silver with purple
venation ; upper lip white.

Measurements in mm.: Snout to vent, 20; width of head, 7; arm,
13; leg, 35.5; axilla to groin, 10.2.

Variation: The five paratypes are all males and agree with the
type in essential details, such as in the constriction on sides of head
behind the tympanum, forward position of nostril, and the narrow
tips of inner toes; the vomerine teeth appear to be present in all
but are completely transparent and scarcely discernible save under
high magnification.

The spine of the pollex is transparent, sharp and in certain cases
does not seem to have pierced the surface of the skin. I presume
that this serves as a "grasping" organ in mating and that the spine
pierces the skin surrounding it during this act. If it is present in
the female it will probably be found to be much reduced. The base
of the first finger lacks nuptual asperities although one can discern
a group of small spherical glands below the skin in the region where
they normally occur in certain other species.

This form stands in relation to Centrolenella much as Plectrophyla
does to Hyla, and probably merits a generic designation. However,
the pollex character may be present but overlooked in certain species

2G2 The University Science Bulletin

referable to the group, and a generic name may already be avail-
able. Until this is ascertained I shall leave it in C entrolenella.

Centrolenella colynibiphyllum sp. nov.

Type: University of Kansas Museum of Natural History, no.
23812 collected at the American Cinchona Plantation, elev. 5,600
ft., Caribbean drainage of Volcan Poas, Costa Rica, July 29.
Richard C. Taylor and E. H. Taylor collectors.

Paratypes: Nos. RCT. 790-791. Same data as type.

Diagnosis: Tympanum moderately distinct, its diameter in dia-
meter of eye at least 2.5 times; width of an eyelid in narrowest in-
terorbital width 1.25 times; frontal region shallowly concave, the
areas about nostrils strongly swollen with a depression between;
canthus indistinct, concave; skin below anal opening forming ver-
tical folds extending under thigh; tibiotarsal articulation 2 mm.
beyond snout; terminal disks of hand wider than digits, fingers less
than one half webbed; choanae large, the diameter of one contained
in distance between them slightly less than two times; no trace of
humeral process; no free pollex rudiment; vomerine teeth absent.

Description of type: Adult male; vocal sac forms ample folds on
chin and throat. Head a little broader than the body ; eyes promi-
nent, directed forward; rounded outline of jaw seen from above in-
terrupted by snout which extends a little farther forward than edge
of mouth; interorbital width small, about one fourth greater than
width of an eyelid, slightly greater than distance between eye and
nostril; frontal region slightly concave; the canthus rostralis in-
distinct, strongly concave; lores oblique, not or but very slightly
concave; nostril distinctly nearer median edge of lip than eye; a
distinct depression between the swollen areas of nostrils; eye pur-
plish black, the color showing through eyelids so they appear dark.
Lower part of tympanum distinct, largely directed upward but ap-
parently covered with thin slightly pigmented skin, the inner edge
more heavily covered.

Tongue subcircular, more or less emarginate on posterior edge,
free only for about one sixth of its length; openings to vocal sac
elongate slits extending along sides of tongue and somewhat -be-
hind; choanae very large, their diameter slightly more than half
distance between them; vomerine teeth absent.

Upper part of arm very slender, forearai thickened with a strong
fold on its outer edge, without a thin flap indicated at elbow ; median
palmar tubercle very distinct; inner tubercle rather indistinct, lack-

Taylor: Genera Centrolene and Centrolenella 263

ing nuptial rugosities but with the minute glandules visible under
the skin; toes broad, terminal disks distinctly wider than digits,
the width of outer finger disk equal to half eye diameter; outer
fingers little more than one third webbed; a trace of web between
first two fingers. Leg elongate, the tibiotarsal articulation reaching
2 mm. beyond tip of snout; heels barely touch when legs are folded
at right angles to the body; a flat rather large inner metatarsal
tubercle; outer tubercle if present very indistinct; toes nearly three
fourths webbed except that first two have webs extending scarcely
half their length.

Skin not granular, but under lens some minute elevations visible,
each with a medial depression suggesting a crater; chin and throat
smooth; abdomen areolate; a very small areolate area on under
surface of thigh; skin below anus forming vertical folds, noticeable
when femurs are at right angles to body.

Color: In life, uniform bright yellow green on back and sides,
with dim suggestions of cream flecks; on ventral surfaces whitish,
semitransparent; tips of toes yellowish. In preservative (formalin
transferred to alcohol), nearly white with numerous star-shaped
chromatophores and indications of small rounded areas without pig-
ment; under surfaces flesh-white without pigment; a very narrow
line of chromatophores on dorsal surface of thigh.

Measurements in mm.: Snout to vent, 26; head length, 7.8; arm,
18; leg, 43.

Variation: The two paratypes taken at the same time and place
differ negligibly, save that the tympanum is a little smaller and a
little less distinct, and the tibiotarsal articulation reaches only to
the tip of the snout. The measurement variations are given in the
following table:

Centrolenella colymbiphyllum sp. nov.

No.

Total
length

Head
length

Head
width

Arm

Hand

and

fingers

Leg

Foot
and
toe

Tibia

Femur

791

26

7.8

9.5

18

8

46.2

20

15

13.7

790

25.8

8

9.2

18

8

43

18.8

13

12.8

23812

25

7.4

9

16.5

7.7

44

19

13.8

]3.5

Remarks: Centrolenella albomaculata differs from this form in
having the snout less protruding, the hand more than half webbed,
the feet four fifths webbed, the vomerine teeth present, and the
nostril closer to the eye. The dorsal color is lavender dotted with
numerous cream spots. The eyelid is purplish black.

Centrolenella granulosa differs in having the dorsum strongly
granular, the nostrils closer to the eye than middle edge of lip, the

264 The University Science Bulletin

snout sloping obliquely in front of the nostrils, the vomerine teeth
present and the choanae much farther apart. The webbing on hand
and foot is distinctly greater.

Centrolenella valerioi differs in having a very small tympanum
(one sixth the eye diameter) , and the interorbital space twice width
of an eyelid.

C. fleischmanni differs in having the eyelid golden, the under
surface of the thigh very strongly areolate on the entire under sur-
face, a distinct outer metatarsal tubercle present, choanae farther
apart, and the frontal area not concave. Both species have the vom-
erine teeth absent and the degree of webbing about the same.

The character of the pollex in C. spinosa distinguishes that form
from all other members of the genus.

The presence of vomerine teeth, numerous white punctations
below the eye and the granulation of the underside of the thigh
distinguishes C. pulveratum.

The specimens were collected at night from leaves of plants grow-
ing in a small mountain stream. They appeared to be very wary
and numerous individuals dived into the water and escaped. In
the same identical habitat a series of Centrolene prosoblepon were
taken. These were apparently less wary and more easily captured.

There is a vague possibility that C. colymbiphyllum is identical
with Boulenger's Hyella puncticrus that has been synonymized with
Centrolene pi^oseblepon by Neiden (1923).

Centrolenella jieischnianni (Boettger)

Hylella fleischmanni Boettger, Ber. Senck. Naturf. Gesells., 1893, pp. 251-253. (Type lo-
cality, San Jose, Costa Rica.)

Centrolenella jleisclunanni Noble, Proc. Biol. Soc. Wash., vol. 37, 1924, p. 69 (combination
suggested).

Three typical specimens are from the environs of Cartago, Costa
Rica. They were taken from low branches above flowing water on
a rainy evening.

Centrolenella valerioi (Dunn)

Centrolene valerioi Dunn, Occ. Papers Boston Soc. Nat. Hist., vol. 5, 1931, pp. 397-399.
(Type locality, La Palina, Costa Rica.)

A single specimen was obtained at El General, Costa Rica, cling-
ing to a leaf of a plant growing in the mire about a small spring.
The dorsal color in life was green with some slightly reticulated
lighter green markings. The eye was golden.

In preservation the specimen is nearly white, peppered with fine
purplish flecks; the area above the eye is cream white but bearing
purplish flecks; the pupil is gray and lavender.

Taylor: Genera Centrolene and Centrolenella 2G5

The specimen measures 20 mm. snout to vent; width of head, 7.5
mm.; arm, 13 mm.; leg, 32 mm.

The tympanum is visible but covered with skin and measures ap-
proximately 1/5 of the diameter of the eye.

It bears considerable resemblance to Centrolenella jieischmanni,
but the arms and legs are proportionately longer and the tympanum
is present.

Centrolenella granulosa sp. nov.

Type: R. C. T. no. 2463. Collected at Los Diamantes one mile
south of Guapiles, Costa Rica, Sept. 7, 1948, by Richard C. Taylor.

Paratype: University of Kansas no. 23802. Collected with the
type; same collector.

Diagnosis: A rather large species of the genus (28 mm.) with
strongly granular skin on head, back and sides of body, dorsal part
of limbs, venter and undersurface of thighs. Large vocal sac, its
surface rugose. Web on outer fingers extending beyond level of
the two outer subarticular tubercles; between second and third
fingers, 1/5 webbed with broad fringe on outer side of second finger;
scarcely a trace of web between first and second; nuptual asperities
on thumb; large inner, very small outer metatarsal tubercle; no
projecting rudiment of a pollex; tibiotarsal articulation 2 mm. be-
yond tip of snout; nostrils equidistant from eye and median tip of
snout; eyelid purple, body (in alcohol) creamy white with numerous
scattered purple or lavender spots.

Description of type: Head broader than body, the eyes strongly
protruding, directed somewhat forward, the pupil horizontal; can-
thus rostralis distinct, slightly rounded; area about nostrils some-
what elevated with a slight depression between them ; in front of nos-
tril, snout slopes forward and down to edge of lip; loreal region slop-
ing more gradually to lip; interorbital distance about 1 to 11/2 times
width of eyelid; tympanum present, its diameter practically three
times in longitudinal diameter of eye; a supratympanic fold from
eye to near insertion of arm.

Tongue cordiform, slightly notched behind; opening to vocal sacs
are curved slits reaching from level of middle of tongue to consid-
erable distance behind tongue ; choanae large, circular, the diameter
of one contained in distance between them four times; two very
indistinct vomerine ridges between choanae, each bearing two or
three vomerine teeth.

266 The University Science Bulletin

Fingers short and broad, the tips dilated, that of fourth finger one
half diameter of eye; a mere trace of web between first and second
fingers; about one fourth to one fifth webbed between second and
third but a rather distinct fringe reaches to terminal pad on outer
side of the second ; between third and fourth a little more than two
thirds webbed, a fringe reaching terminal pad on outer side of third
finger; subarticular tubercles well defined; a large, very distinct
oval palmar tubercle; large tubercle at base of first finger bearing
colorless nuptual asperities which extend much beyond tubercle and
across the dorsal surface; a strongly defined fold on underside of
forearm.

Legs long, the heels overlapping little more than two millimet-
ers; tibiotarsal articulation extends beyond tip of snout about two
or three millimeters; toes three fourths to four fifths webbed; sub-
articular tubercles distinct, rather large; a strong inner metatarsal
tubercle; and a very small outer; a very slight, scarcely distin-
guishable inner tarsal fold.

Granulations on skin strongly defined on back, sides, head, and
to a lesser extent on surface of forearm and lower part of leg and
foot; on throat and chin, skin rugose but apparently not granular;
venter and the greater part of underside of thigh, strongly areolar
or granular, the granules being very much larger than those on
back; those under the thigh distinctly larger than those on venter;
a narrow, but relatively long transverse free flap covers anal opening.

Color: In life, leaf green with some slightly darker areas evi-
dent; below transparent white or flesh; in preservative (formahn
transferred to alcohol) color cream flesh with a very faint wash of
lavender with numerous (about 40) small purple spots; under a
lens the lavender wash is seen to be caused by very numerous, small
chromatophores. All ventral surfaces, and much of the lateral sur-
faces of limbs and body, cream- white lacking pigment; eyelids
purple; eyes silver with some purplish reticulation.

Measurements in mm.: Snout to vent, 28.2; width of head, 10;
axilla to groin, 16; arm, 18; leg, 49.

Variation: The paratype is somewhat more lavender than the
type. The dorsal spots are smaller and more numerous on the back
and on the tibia. The iris of the eye is darker; the vomerine tooth
patches are more distinct. Otherwise the two agree in all essential
details.

Remarks: The absence of a humeral hook or process places this
species in the genus Centrolenella Noble. Noble regarded the ab-

Taylor: Genera Centrolene and Centrolenella 267

sence of vomerine teeth a character of the genus. However the
teeth are not invariably present. The teeth may be absent in some
individuals of a species and present in others.

The position of the nostril, halfway between eye and the median
tip of snout, the purple eyelid, the very granular skin, the longer
leg, presence of a tympanum, absence of a pollex rudiment, and
the markings, will together distinguish this form from other species
of Centrolenella.

The specimens were taken at night on small trees growing on the
edge of a small river.

Centrolenella albomaculata sp. nov.

Type: University of Kansas Museum of Natural History no.
23814. Los Diamantes, one mile south of Guapiles, Costa Rica,
Sept. 7, 1947, E. H. Taylor and Richard C. Taylor, collectors.

Paratype: R. C. T. no. 2020. Same date, locality, and collectors.

Diagnosis: A rather large sized species of the genus (27 mm.
snout to vent) ; tympanum distinct, nostrils slightly closer to eye
than to median point of snout at tip ; interorbital distance equal to
width of eyelid ; heels overlapping ; tibiotarsal articulation 4-5 mm.
beyond tip of snout; toes five sixths webbed, the membrane touch-
ing the terminal expansions on one side or other of all toes, save
fourth; a distinct web present at base of first and second fingers;
posterior part of back slightly but indistinctly granular; vomerine
teeth strongly developed in two ridges near posterior level of cho-
anae. No pollex rudiment; purplish lavender with numerous cream
spots on all unconcealed dorsal surfaces.

Description of type: Head a little wider than body, the eyes
large, protruding, their diameter equal to their distance from tip
of snout; areas about nostrils strongly swollen, with a strong de-
pression between and somewhat behind them; canthus rostralis
wanting; in front of nostrils the snout slopes to the edge of lip at
an angle distinctly greater than slope of loreal region; lores dis-
tinctly concave; tympanum small, its greatest diameter contained
in distance from eye one and three fourths times, and in the longi-
tudinal diameter of eye a little more than three times.

Tongue rather rounded, not or but very slightly emarginate pos-
teriorly, free for about one fifth its length; opening of vocal sacs
elongate, curved slits lying partly at side of tongue and extending
some distance behind; choanae large, their diameter contained in
space between about three times; vomerine teeth in two raised

268 The University Science Bulletin

patches lying between the posterior level of choanae and extending
somewhat behind this level; (in paratype not reaching posterior
level of choanae).

Digits short, widened at the tips, that of third finger a little more
than one half eye diameter; a very narrow web between first two
fingers; second and third fingers one third webbed, the web extend-
ing as a fringe to the terminal expansion on outer side of second
finger; third and fourth toes two thirds webbed, the web extending
beyond level of subarticular tubercles; latter strongly developed;
an elongate pad at base of first finger and very distinct palmar pad
at base of wrist ; base of first finger with an area of nuptial asperi-
ties; supernumerary tubercles indistinct; upper arm slender, fore-
arm thickened with a well-developed fold on under side; legs long
and slender, the tibiotarsal articulation extending several milli-
meters beyond tip of snout when leg is adpressed to body; heels
overlap 2.2 mm.; toes four fifths webbed, the web reaching the
terminal pad of each toe on one side, except fourth; terminal pads
truncate, that of inner toe rather pointed; a rather large inner
metatarsal tubercle, rounded anteriorly, the tip slightly free ; outer,
if present, very indistinct; subarticular tubercles well developed;
some distinct supernumerary tubercles; inner tarsal fold scarcely
indicated.

Skin with some indistinct granulation on posterior part of back;
slightly rugose anteriorly and on sides of body; upper surface of
limbs also slightly rugose; throat and breast smooth; abdomen
granular as is the greater part of under side of thigh; an enlarged
granule on each thigh medially ; segmentation of abdominal muscles
distinctly indicated on venter; a small free flap over anal opening.

Color in life: Yellow green on all exposed surfaces, with indi-
cations of small cream dots; all concealed surfaces creamy white
nearly transparent; in preservative (formalin transferred to al-
cohol) purplish to dark lavender above, somewhat lighter on the
upper surface of limbs with very numerous rounded cream spots or
punctations, somewhat larger on hind limb than on back; venter
and under surface of limbs cream white; eyelids dark purple with
white flecks ; iris purple with minute silver flecks ; some white flecks
in tympanic and anal regions.

Measurements in mm.: Snout to vent, 26; width of head, 9;
arm, 19; length of hind limb, 50; axilla to groin, 14.

Variation: The single paratype collected in the same locality
with the type agrees in all essential details save that the vomerine

Taylor: Genera Centrolene and Centrolenella 269

teeth are somewhat farther forward; and outer metatarsal tubercle
is indicated. The color has been somewhat changed due to rust in
the preserving fluid.

A key to the recognized species (for preserved material) is here
given :

1. Humeral hook in males; color lavender with numerous small purple spots and
minute chromatophores; nostrils far forward, nearer median edge of lip than eye;
tympanum distinct; type 24.5 ^ ; greatest length, 27.5 5 .. .Centrolene prosoblepon
No humeral hook in males 2

2. A rudiment of a pollex present, free at tip, terminating in a sharp firm spine in
males, condition in females not known ; skin of back and sides under lens minutely
rugose but not granular ; snout nearly vertical in front, nostrils nearer median
point of lip than eye; lavender with neither light nor dark spots (females not
known); vomerine teeth present; length, 21 mm. ^ .. .Centrolenella spinosa sp. nov.
No free rudiment of pollex bearing spine 3

3. Tympanum concealed; vomerine teeth absent; nostrils not elevated; color cream-
flesh, unspotted but with numerous lavender or purple chromatophores visible
under lens; upper eyelid golden yellow or creamy yellow .. .Centrolenella fleischnianni
Tympanum not concealed ; vomerine teeth present or absent 4

4. Tympanum minute, one sixth of eye diameter or "very small" 5

Tympanum larger, at least one-third of eye diameter 6

5. No vomerine teeth present ; first finger longer than second ; canthus distinct but
rounded ; outer fingers about one-half webbed ; interorbital space twice width of
an eyelid ; heel reaches beyond snout ; skin smooth above ; white with a few faint
dark [purplish?] chromatophores; type, 21 mm.; sex not mentioned. (Said to

reach 26 mm.) Centrolenella valerioi

Tympanum very small covered with pigmented skin ; vomerine teeth present ; back
smooth ; first and second fingers equally long ; outer fingers about y-i webbed ;
toes except fourth nearly completely webbed ; colorless with scattered small punc-
tations, numerous on the side of head Centrolenella pulveratum

6. Vomerine teeth absent; skin lacking granules on back; nostril much nearer
median edge of lip than eye ; interorbital space one fourth wider than an eyelid ;
outer fingers less than half webbed ; diameter of choanae in distance between

choanae minutely less than two times Centrolenella colymbiphyllum

Vomerine teeth present; dorsal skin more or less granular; nostril nearer eye than
median edge of lip 7

7. Skin strongly granular above; nostrils nearer eye than tip of snout; tip of snout
oblique, not vertical ; vomerine teeth weakly developed ; white or slightly lavender
with numerous purple spots, and finely scattered lavender chromatophores; outer
fingers two thirds webbed ; ejelid deep purple ; 28 mm. ^

Centrolenella granulosa sp. nov.
Skin weakly granular on posterior part of back, somewhat rugose anteriorly and
en head; vomerine teeth strongly developed; tibiotar.sal articulation 4-5 mm.
beyond snout; interorbital width equal to width of eyelid; diameter of tympanum
in eye diameter a little more than three times ; nostril minutely nearer eye than
lip ; lavender above, with numerous cream spots on all exposed surfaces ; type,
27 mm Centrolenella albomaculata sp. nov.

270 The University Science Bulletin

LITERATURE CITED

1892. BoETTGER, Oscar, Katalog der Batrachier — Sammlung im Museum der

Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am
Main. Aug., 1892, pp. i-x; 1-73.

1893. Ein neuer Laubfrosch aus Costa Rica. Ber. Naturf. Gesells.,

Frankfurt am Main. 1892-1893 (1893), pp. 251-252.

1896. BouLENGER, G. A., De.scription.s of new Batrachians collected by Mr.

C. F. Underwood in Costa Rica. Ann. Mag. Nat. Hist., ser. 6, vol. 18,

1896, pp. 340-342.
1931a. Dunn, Emmet R., New Frogs from Panama and Costa Rica. Occ.

Papers Boston Soc. Nat. Hist., vol. 5, Aug. 18, 1931, pp. 385-401.
1931b. — - — The amphibians of Ban'o Colorado Island. Occ. Papers Boston

Soc. Nat. Hist., vol. 5, Oct. 10, 1931, pp. 403-421.
1933. Amphibians and reptiles from El Valle de Anton, Panama. Occ.

Papers Boston Soc. Nat. Hist., vol. 8, June 7, 1933, pp. 65-79.

1923. NiEDEN, Fritz, Das Tierreich. 46 Lief. Anura I. 1923, June, pp. i-

xxxii -f- 1-584, figs. 1-380. Berlin and Leipzig, Walter de Gruyter and
Co.

1924. Noble, G. K., Some neotropical batrachians preserved in the United

States National Museum with a note on the secondary sexual char-
acters of these and other amphibians. Proc. Biol. Soc. Washington,
vol. 37, Feb. 21, 1924, pp. 65-72.

1925. An Outline of the Relation of Ontogeny to Phylogeny within the

Amphibia I. American Mus. Novit., no. 165, Apr. 16, 1925, pp. 1-17.

1873. Peters, W., iiber eine neue Schildkrotenart, Cinostemon Effeldtii und
einige andere neue oder weniger bekannte Amphibien. Mon. Akad.
Wiss. Berlin, 1873, pp. 603-618, pi. 5.

I

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. I] April 20, 1949 [No. 5

Two New Teiid Lizards from Costa Rica

BY

Edward H. Tatloh

Abstract. — Two species of lizards belonging to the Family Teiidae are de-
scribed as new. One species, Alopoglossus -plicatus, is the first record of that
genus on the North American continent. The second species, Leposoma bi-
secta, is the second species of that genus known north of the Isthmus of
Panama.

The lizard family Teiidae, which apparently has undergone the
greater part of its evolution in South America, is represented there
by about 28 known genera, a number greater than is possessed by
any other South American lacertilian family at the present time.
Eight genera have entered the isthmus and have pushed various
distances to the north. These are Cemidoj)horus, a genus wide-
spread in Central America, Mexico and the United States; Ameiva,
a genus with several representatives in Central America and Mexico
extending far to the north in eastern Mexico; a third, Gymnoph-
thalmus, reaching at least to the isthmus of Tehuantepec ; Leposoma
and Anadia, known to have reached as far north as Costa Rica, and
Echinosaurus and Ptychoglossus known in Panama.

A species encountered in the east slope drainage of Costa Rica
at an elevation of about 2,600 feet is referred to the genus Alopo-
glossus and is here described as new. There may be some doubt
about the use of Alopoglossus for those lizards having chevron-
shaped plicae on the tongue instead of scales ; Boulenger recognized
the genus Alopoglossus for these forms, but Burt and Burt (1931)
synonymized it with Pantodactyhis Dumeril and Bibron. The dif-
ferences apparently are of generic significance.

(271)

272 The University Science Bulletin

Alopoglossus pUcatus sp. nov.

Figure 1.

Type: University of Kansas Museum of Natural History, no.
23800. Taken on the Morehead Finca, 5 miles southwest of Tur-
rialba, Costa Rica, July 21, 1947, by Edward H. Taylor.*

Diagnosis: Dorsal scales around body forming nearly regular
transverse and oblique keeled series ; 30 scales from occiput to base
of tail; 24 scales from chinshields to anals; 33-35 scales around
body at middle, all strongly keeled save on belly ; keels terminating
in spines, the spine reduced or absent on lateral rows; ventrals in
four regular, longitudinal, smooth, quadrangular rows, outer larger
than inner; gular collar indicated; four supraoculars; venter cream
with dark spots or flecks in males (female may be white below).

Description of the type: Rostral nearly twice as wide as high
forming a straight (or very slightly curved) suture with the fron-
tonasal; latter large, in contact laterally only with nasal; prefrontals
pentagonal, touching only preocular and first supraocular laterally;
frontal wider near anterior end than at posterior end, sides slightly
concave, ends broadly angular, in contact with three supraoculars;
two frontoparietals, larger than prefrontals but at least one fourth
smaller than frontal; parietals large, longer than wide, much larger
than other head scales, angular at anterior end, somewhat rounded
posteriorly, and separated from each other by a large parallel-sided
interparietal.

Nasal large, triangular, at least partially divided by a suture
running up from nostril; a rectangular preocular much higher than
wide; four supraoculars, four superciliaries, anterior largest; 2 rows
of ciliaries on upper eyelid; lower eyelid with a more or less trans-
parent disk divided into three parts; 2 presuboculars precede an
elongate curved subocular; 3 presuboculars, the upper touching the
anterior superciliary; a small "loreal."

Seven upper labials, third greatly elongated, lying for the most
part below eye; mental deeper and wider than rostral; six lower
labials; large azygous postmental followed by three pairs of chin-
shields, first and second pairs in contact mesially and in contact
with labials laterally, third pair separated mesially from each other
and from labials laterally, followed by an enlarged pair of scales
separated mesially; six transverse rows of smooth scales between

* TTie University of Kansas Natural History Museum wishes to acknowledge financial aid
to the Endownaent Association of the University of Kansas, in the making of collections in
Costa Rica.

Taylor: New Teiid Lizards

273

this last pair of scales and the groove formed by gular collar; six
rows of temporal scales, posterior scale of upper row largest, some-
what ridged; some slight ridges on some of the other temporal
scales; auricular opening large.

Fig. 1. Alopoglossus plicatus sp. nov. Univ. Kansas Nat. Hist. Mus.
no. 23800 Type. Five miles southwest Turrialba, Costa Rica. A. Head
lateral view. B. Head ventral view. C. Head dorsal view. (Slightly
diagrammatic.) Snout to ear length 11.6 mm.

Dorsal scales strongly keeled, their sides more or less parallel,
terminating in a spine, continuing to tip of tail; scales forming
transverse rows. Lateral scales on neck small, smooth, irregular in
size ; around arm insertion a rather large granular area present, and
a similar one of lesser extent about leg insertion.
18—3659

274 The University Science Bulletin

Ventral scales of breast and abdomen large, quadrangular,
smooth, beginning at collar where scales are slightly curved, and
continuing back to level of thigh, arranged in four longitudinal
rows and sixteen transverse rows, the outer larger than inner;
outer row on each side bordered by a series of somewhat enlarged
scales about one half as wide ; scales under tail smooth ; four elon-
gate anal scales, outer smaller than inner, separated from the last
modified row of the abdominal series by a row of small scales, two
on each side, bearing femoral pores; total series of femoral and
preanal pores 12 (13) on each side.

Arms and legs moderately developed, pentadactyl; digits well
developed and clawed; anterior surface of arm with large, smooth,
irregularly-shaped scales; those on back and ventral surface much
smaller; front scales on thigh smooth and large; dorsal scales on
thigh smaller, strongly keeled; posterior thigh surface covered with
granular scales; under surface of tibia with large smooth scales;
those on other surfaces smaller, keeled; two enlarged scales, each
with a sharp cutting edge, at base of first finger.

Each side of tongue decorated with a regular series of plicae that
run forward to medial line, forming 24 chevrons; posteriorly there
are eight or nine rows on each side that fail to meet in the middle.

Measurements (in mm.): Snout to vent, 50; snout to arm, 19.2;
axilla to groin, 25; head width, 8.4; snout to ear, 11.6; tail, 60 (re-
generated); arm, 13; leg, 19.5.

Color in life: Head nearly uniform mahogany brown, slightly
darker in supraorbital region; back slightly reddish-brown with
numerous dark flecks; a dorsolateral light line from above tympa-
num to behind level of arm insertion; sides blackish, but with very
numerous cream-white flecks, some suggesting lines; venter cream
with numerous black spots or flecks; black marks on the ventral
scales are largely on scale edges and these, being contiguous, tend
to form irregular black stripes. Chin with rounded dark spots.

Tail partly regenerated, proximal part colored much like body;
ventrally, dark spotting tends to form transverse rows of dots or
narrow irregular bars; regenerated portion of tail much lighter with
a reddish cast; only a few minute dark flecks on sides and a very
few on underside. There is somewhat more black mixed in dorsal
coloration of body in area between the two anterior dorsal light
lines, this area being considerably darker than elsewhere.

Relationship: This species is probably most closely related to

Taylor: New Teiid Lizards 275

Alopoglossus buckleyi. From the species which Boulenger placed
in this genus, A. plicatus may be distinguished as follows:

From Alopoglossus copii by the presence of quadrangular scales
on the neck, rather than the more typical lanceolate triangular,
spine tipped scales of copii. There are, of course, other differences
including a longer series of femoral pores.

From carinicaudata it differs in having 33 - 35 instead of 21 scales
round the body; and from buckleyi in having smooth instead of
keeled scales, more scales around body, and a very different color-
ation.

Variation: The paratype, a young female from the same locality,
differs somewhat from the type, but I suspect that this is due largely
to age and sex. The lateral groove on the tail is veiy distinct, and
that on the dorsal surface of the tail is a little less distinct than
the lateral. The femoral pores are absent and there is a distinct
difference in the character of the scales that precede the anal scales.
These are proportionately much larger, while the preanal scales are
distinctly smaller than in the type.

In this paratype, there are 34 scale rows around the middle of
the body and 27 between the chinshield and the anal opening. The
scales of the row bordering the outer of the four median ventral
scale rows is distinctly larger than in the type and one might well
count six ventral rows instead of four.

The ventral surface of tail and body are immaculate cream; the
chinscales have a few black spots. It is possible that adult females
may develop more pigment on the venter. On the other hand it is
not impossible that a second form is involved. Only larger series
can determine this beyond question.

Leposoma bisecta sp. nov.

Type: University of Kansas Natural History Museum, no. 23801,
collected on the Pacific slope, i/^ mile west of El General, Costa
Rica, at an elevation of about 300 meters, Aug. 21, 1947, by Edward
H. Taylor.

Diagnosis: A species characterized by a large divided fronto-
nasal, narrowed anteriorly; the scales of head strongly striate longi-
tudinally; ventral scales and gular scales all very strongly keeled;
four supraoculars ; prefrontals short, as wide as long ; no postfrontal ;
four preoculars; a pair of frontoparietals; interparietal larger than
parietals. Dorsal scales large, more than three fourths as broad
as long, heavily keeled and with a short posterior spine, forming

276

The University Science Bulletin

transverse and oblique rows on back and sides. No trace of a gular
collar.

Description of the type: Rostral slightly more than twice as wide
as high; frontonasal divided into two elongate scales narrowing an-
teriorly; a pair of prefrontals nearly as broad as long, in contact
with each other for a little more than half their length (due to an
injury one is partially fused to frontal) ; frontal concave laterally,
broadly angular anteriorly and posteriorly, one and one third times

B

Fig. 2. Leposoma bisecta sp. nov. Univ. Kansas Nat. Hist. Mus. no.
23801, Type. El General, Costa Rica. A. Head dorsal view. B. Head
ventral view. Actual head width 6.7 mm. (Slightly diagrammatic.)

as long as wide; frontoparietals pentagonal, in contact with each
other for half their length; interparietal larger than the parietals
and separating them. Scales following the parietals and inter-
parietal not or but little differentiated from other neck scales; four
supraoculars. All dorsal head scales with either two, three or four
heavy ridges or keels.

Nostril pierced in a large nasal, which has a slight groove above
nostril, suggesting a suture; five scales in combined subocular-pre-
ocular series, including "loreal"; four superciliaries; two or three
small postoculars not distinguished from other temporal scales;

Taylor: New Teiid Lizards 277

lower eyelid with a transparent disk divided into two parts; seven
upper labials in following ascending order of size: 6, 1, 5, 4, 3, 2, 7;
five lower labials ; a large azygous postmental followed by two pairs
of chinshields which are in contact mesially and border the labials
either entirely or partially (second pair) ; a third pair much smaller,
separated from the labials by two or three scales, and from each
eath other by a slender elongate scale; each of the third pair fol-
lowed by three scales, these separated by two or three smaller
scales. There follows a row of very small keeled scales that ex-
tends to the very large auricular opening. Seven rows of temporal
scales; postauricular scales pyramidal or subconical. No trace of a
gular collar.

Scales of dorsum large, imbricate, heavily keeled, each terminat-
ing posteriorly in a small spine, forming somewhat irregular trans-
verse series and regular oblique series; 26 scales in row about middle
of body; 32 scales from occiput to posterior level of thighs; 35 rows
from chinshields to preanal pore-scales; four contiguous preanal
pore-scales immediately preceding preanal scales. Scales of breast
and abdomen arranged in regular transverse series in six longitu-
dinal rows, all keeled strongly, the keel terminating in a spine pos-
teriorly. These scales differ from gular series in having their sides
more nearly parallel. Five preanal scales border the vent, arranged
as follows, a small median preceded by a larger triangular scale;
these two bordered laterally by two elongate widened scales, nar-
rowed at each end; outermost scales small. Femoral pores 7-7,
reaching to the continuous preanal pore series of four (2-2),

Limbs pentadactyl, all clawed, inner finger and toe reduced;
scales on upper surfaces of arm more or less regular, keeled; those
on under surface smaller, some tetrahedral with keels reduced;
scales about arm insertion small, more or less tubercular; scales
on front and dorsal surface of legs irregular in size, more or less
similar in shape; scales on posterior thigh granular with a patch of
slightly larger scales; scales in groin, and about leg insertion,
smaller.

Tail, regenerated for the most part, somewhat compressed, bear-
ing a more or less distinct lateral groove; scales on regenerated
part of tail similar to those of original tail.

Auricular opening very large, nearly equal to eye opening.

Tongue covered with scales; no suggestion of lingual plicae or
chevrons.

Color: Dark indefinite brown, the scales with minute black flecks

278 The University Science Bulletin

or punctations; a light, tan, dorsolateral line from middle of neck
to some distance on tail, distinct anteriorly, less distinct in middle
of body, forming a series of light dots on tail; sides of neck and
body darker; head blackish on top and sides; venter light, immacu-
late, save for a few flecks under tail; regenerated part of tail darker
above and below; white spots on upper and lower labials alternate
with similar dark areas; a few cream spots on side of neck. Scales
or lamellae under feet and toes blackish.

Measurements (in mm.): Snout to vent, 34.5; snout to arm,
13.2; axilla to groin, 17.3; tail (regenerated), 43; arm, 10.2; leg,
13; head width, 6.7; head to auricular opening, 8; head length, 10.

Relationships : I am associating this species with the genus Lepo-
soma despite the fact that the frontonasal is paired. On the basis
of this character it does not appear to be closely related to other
members of the genus.

Remarks: Burt and Burt (1931) give meagre data on two speci-
mens of a Leposoma from Suretka, Costa Rica (M. C. Z. nos. 18916-
18917), which they refer to the species Leposoma. dispar Peters.
They record certain differences that obtain between the specimens
and the type of dispar, and it is not impossible that their specimens
warrant description under a different name. As far as data is avail-
able, the specimens mentioned differ from the type here described
as follows:

Specimens Type Specimen of L. bisectn

Suretka, Costa Rica El General, Costa Rica

1. Head shields roughened poster- 1. All head shields with one to three
iorly. strong keels.

2. Twenty - seven to twenty - eight 2. Thirty-two scales from occiput to
scales from occiput to base of tail. base of tail.

3. Nineteen to twenty scales about 3. Twenty-six scales about middle of
middle of body. body.

4. Thirty to thirty-two scales from 4. Thirty-five scales from chinshields
chinshields to anals. to anals.

Apparently they agree in having four supraoculars. They differ,
however, in the character of the' frontonasal and there may be other
unstated differences.

LITERATURE CITED

1931. Burt, Charles E., and May Danheim Burt. South American Lizards
in the collection of the American Museum of Natural History. Bull.
American Mus. Nat. Hist., vol. 61, 1931, pp. 227-395, figs. 1-15.

THE UNIVEESITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. I] April 20, 1949 [No. 6

New Salamanders from Costa Rica

BY

Edward H. Taylor

Abstracjt. — A short review of the known species of salamanders of Costa
Rica is given, and four new species are described. These are Magnadigita
nigrescens, Bolitoglossa palustris, Parvimolge richardi and Oedipina serpens.

The known salamander fauna of Costa Rica consists of some
ten plethodontid species representing several genera. One other
species, Oedipina coUaris, has been reported both in Nicaragua
(its type locality) and Panama, and it may be expected in Costa
Rica if its range is continuous.

The following species have been taken, or at least reported in
Costa Rica:

Pseudoeurycea? picadoi Magnadigita robusta

f Bolitoglossa platydactyla Magnadigita subpalmata

Bolitoglossa lignicolor Oedipina unijormis

Bolitoglossa striatula Oedipina alfaroi

fBolitoglossa flaviventris Haptoglossa pressicaucla

Cope (1876) reported a specimen of Oedipus moriof "a partly
preserved specimen from the eastern slope of the Pico Blanco."
This has been referred to the synonymy of Oedipus subpalmatus
{^Magnadigita subpalmata] by Dunn (1926). Again Cope (1887)
lists Oedipus morio and includes with the above reference a report
of a specimen collected by Zeledon on the "Plateau of Costa Rica."

Giinther (1902) reports a species which he designates as Spel-
erpes morio from the plateau of Cartago (Zeledon collector), and
seemingly refers to the same material mentioned above. Dunn
(1926) has referred this reference to the synonymy of what Dunn
designates as Oedipus morio (corrected by Schmidt, 1933) to Oedi-

(279)

280 The University Science Bulletin

pus dunni. Since the reference also contains specimens from Mex-
ico and Guatemala it is not certain whether one may accept the
species dunni as a part of the Costa E,ican fauna or not.

Bolitoglossa platydactyla {Spelerpes variegatus) is reported from
San Carlos, Costa Rica, by Giinther (1902). Dunn (1926) states
that he has examined this species and that he identifies it as being
Oedipus platydactyla. He also identifies two British Museum speci-
mens from Medellin, Colombia, as 0. platydactyla. If they are cor-
rectly identified the extent of the range (from Eastern San Luis
Potosi, Mexico, to Medellin, Colombia) is greater than that of any
other Central American salamander. He does suggest the possibility
that these specimens "may represent an undescribed form." Cope
reports the species as Oedipus variegatus from Buenos Ayres in
southwestern Costa Rica.

A specimen purporting to be from Miravalles, Costa Rica, was
referred by Dunn (1926) to Oedipus salvinii. Schmidt (1936) has
shown that Dunn has confused Oedipus flauiventins Schmidt with
Oedipus salvinii and places Oedipus salvini Dunn (part) in the
synonymy of the latter. The Costa Rican specimen agrees with
flaviventris in having the venter "light." Schmidt does not make
specific mention of the Costa Rican specimen, but I suspect it should
be regarded as flaviventris until proved to the contrary. The re-
maining species of the list seem to be unquestionably a part of the
Costa Rican fauna.

Pseudoeuryceaf picadoi (Stejneger). The type of this species is
from La Estrella, southeast of Cartago, Costa Rica. Specimens
have been taken at La Raima, and certain of the type series of sub-
palmatus are said to be of this species (Dunn, 1926, p. 380).

Bolitoglossa lignicolor (Dunn) has been reported, by Dunn
(1926), on the basis of a single specimen from Sarapiqui, Costa
Rica. The type locality is Chiriqui, Panama.

Bolitoglossa striatulus (Noble) originally described from Cukra,
near Blufields, Nicaragua, is known from a specimen from Turri-
alba, Costa Rica, and two specimens from Surubres, Costa Rica
(Dunn, 1926).

Magnadigita robusta (Cope) was originally described from the
slopes of Volcan Irazu, Costa Rica. This is a well-known species
and is the largest salamander species occurring in the country.

Magnadigita subpalmatus (Boulenger) was originally described
from La Palma, Costa Rica. It is a well-established form but there

Taylor: Salamanders from Costa Rica 281

i.-. a strong probability that in the literature specimens of other
species have been referred to it erroneously.

Oedipina uniformis (Keferstein) was originally described from
''Costa Rica." Numerous specimens have been reported. Here
again it seems probable that other species have been confused with
it in the literature.

Oedipus alfaroi Dunn, is known from the type, from Zent, and
two specimens from Suretka, Costa Rica.

Haptoglossa pressicauda Cope is known only from the type, which
is now lost.

Giinther, in compiling the Reptilia and Batrachia, of the Biologia
Centrali-Americana, probably had no more than a dozen specimens
from Costa Rica. Dunn, in his "Plethodontidae," lists about 114
specimens from Costa Rica in the material available to him in 1926.
All are placed in the preoccupied genus Oedipus. This material was
distributed as follows:

16 robustus 1 lignicolor

7 picadoi 3 striatula

49 subpalmatus 1 alfaroi

1 flavivcntris (salvinii?) 36 uniformis

Besides these, Dunn examined the following Costa Rican speci-
mens: 1 robustus, 1 subpalmatus and 14 uniformis including the
type of Opheobatrachus vermicularis Gray in the British Museum
of Natural History, and mentions that there were other specimens
not examined (perhaps some of which may have been from Costa
Rica).

In the collection made by the Austrian Biological Expedition of
1930, reported by Wetstein * three species are reported: subpal-
matus, 10 specimens; picadoi, 1; uniformis, 1.

The collections made in the summer of 1947 by Richard C.
Taylor and myself contain representatives of only four of the known
species. No specimens of the reported forms Bolitoglossa platy-
dactyla, B. flaviventris, Pseudoeuryceaf picadoi, Oedipina alfaroi or
Haptoglossa pressicauda, were found. However, numerous speci-
mens of the known Magnadigita robusta, Magnadigita subpalmata,
Bolitoglossa striatula and Oedipina uniformis were taken; several
others were discovered that are believed to represent undescribed
species. A part of these are treated in the following pages.

*Sitz. Akad. Wiss. Wien Abt. 1, Bd. 143, Heft. 1, 2, 1934, pp. 1-39, 9 text figs.

282 The University Science Bulletin

Magnadigita nigrescens sp. nov.

Type: University of Kansas Museum of Natural History no.
23816, collected at Boquete Camp (on highway between Millville
and San Isidro El General), Costa Rica, elevation 6,000 ft., Aug. 17,
1947, by Edward H. Taylor.

Diagnosis: Uniformly blackish brown above, slaty black on sides
and ventral surfaces; hand and foot webbed, the terminal joint of
truncate digits free of web except that of inner digit; thirteen costal
grooves ; nostril small ; paravomerine teeth in a single patch, notched
behind; 11-11 vomerine teeth, arranged partly in two series, be-
ginning outside level of choanae, the two series separated by twice
the diameter of choanae; no sublingual fold; maxillary teeth re-
duced, 7-9.

Description of the type: Adult $ ; head broad, the snout bluntly
rounded, somewhat truncate; a slight swelling on lip below nostril;
eye moderate, its diameter longer than snout ; posterior part of eye-
lids fit under a diagonal fold; a strong gular fold on neck with a
nuchal groove arising from each side, which meets its fellow on the
middorsal line of the neck; anterior to gular fold, a groove com-
pletely crosses chin and ascends either side, crossing behind angle
of mouth and terminating at the postorbital groove ; latter arises at
eye, and extends back but does not reach to nuchal groove; a small
lateral groove in front of the nuchal groove which reaches to the
middorsal line; thirteen costal grooves counting those in axilla and
groin; adpressed limbs separated by between two and one half to
three costal folds; fold of the posterior extension of the hyoid ap-
paratus reaches second costal fold; tail regenerating presumably, a
basal constriction on tail; walls of cloaca with smooth folds; glan-
dular area behind insertion of leg not especially distinct.

Skin generally smooth, but area on snout and interorbital region
somewhat rough ; venter crossed by very numerous fine striations.

Limbs rather short, the very broad digits united by a web, almost
palmate, the terminal joint of the digits, other than the inner, sub-
truncate, with terminal joint free and with a small pad present
below tip; between toes the web may be slightly excised.

Floor of mouth without a free sublingual fold; choanae very
small, their diameter contained in distance between them about six
times ; paravomerine teeth in a single elongate patch widening grad-
ually posteriorly and notched behind, anteriorly narrow and, save
for two or three scattered teeth, widely separated from the vomerine
teeth; the vomerine series of about eleven teeth begin beyond the

Taylor: Salamanders from Costa Rica 283

outer level of the choanae and curve back, separated from each
other by a distance equal to twice diameter of choanae ; each series
arranged partially in two rows; mandibular teeth small, 7-9 almost
covered by gums ; only a single premaxillary tooth present ; 25 man-
dibular teeth on each side of jaw.

The tail (regenerated) is shorter than head and body, with eight-
teen caudal grooves present; width of head in head body length, 5.2
times; length of head to gular fold (lateral) in head body length, 3.9
times.

Color: In life nearly uniform slate black above and below; in
preserved state the dorsal color is dark brown, the sides slate black,
the venter blackish brown ; under side of limbs lighter brown ; under
side of hands and feet grayish white ; chin lighter brown than body.

Measurements {in mm.): Total length (tail regenerated), 70;
snoat to arm, 15; snout to end of vent, 46; axilla to groin, 25; arm,
9.8; leg, 10.1; head width, 8.8; head length to gular fold (lateral),
11.8; head length to gular fold (median), 11.8.

Remarks: Only a single specimen of this species was taken. It
was obtained from within the naturally hollow stalk of a large-
leafed plant. The entrance had been gained through a hole bored by
an insect. It was found nearly a meter below the point of entrance
when the stalk was accidentally split.

The curious reduction of the maxillary teeth occurs in no other spe-
cies of the genus that I have examined and I am uncertain as to its
closest relationship.

Bolitoglossa palustris sp. nov.

Type: University of Kansas Museum of Natural History, no.
23817, collected at San Isidro el General, August 23, 1947, by Ed-
ward H. Taylor.

Diagnosis: A member of the rufescens group, with a short
broad head; fully webbed or palmate hand and foot; vomerine teeth
16-18 on strongly elevated ridges, in more than a single series be-
ginning about level of middle of choanae; choanae large; maxillary
teeth 14-14; premaxillary teeth 4; no free sublingual fold under
tongue; costal grooves indistinct, probably 13; tail much shorter
than head and body.

Description of type: Snout moderately short, truncate, mod-
erately elevated above the mouth; subnarial swellings small dis-
tinct; nostril small; canthus rostralis rounded, the loreal region
sloping abruptly to lip ; diameter of eye greater than length of snout,

284 The University Science Bulletin

about equal to its distance from middle of tip ; surface of snout on a
level with interorbital and occipital surfaces; interorbital width,
equal to twice width of eyelid; upper eyelid not overlapping lower
behind eye, the ends not fitting in a small groove; no orbitolabial
groove; a postorbital groove or depression reaching back to a ver-
tical groove from side of chin which crosses the angle of jaw but
fails to reach the top of head, nor is it continuous across chin; a very
well-defined gular fold, but grooves from ends of fold not or scarcely
traceable on sides of neck, and definitely not reaching to median
nuchal line; angle of mouth reaching slightly behind level of orbit;
posterior extension of hyoid element forming a very strong fold
which reaches back of arm insertion half the length of arm.

Maxillary teeth 14-14 (possibly two or three more as evidenced
by breaks in the continuity of the series) ; 4 premaxillary teeth ; vom-
erine teeth, 16-18; teeth arranged in more than a single irregular
row beginning at the level of middle of choanae, and curving
slightly, separated from its fellow by a distance equal to greatest
diameter of choanae; their diameter contained in distance between
them, three and one half times ; paravomerine teeth in a single sub-
triangular patch, the length only about a third greater than width,
not notched behind, separated from the vomerine series by a dis-
tance greater than half distance between choanae, teeth strong form-
ing typical chevron-shaped series; some pigment evidence in anterior
palatal region at side of paravomerine teeth and on floor of mouth
anterior to tongue.

Dorsal surface of back rather rough ; side and venter smooth ; cos-
tal grooves not very distinct, apparently 13 present; adpressed limbs
separated by 3V2 folds; a constriction at base of tail; 25 caudal
grooves are present ; tail tetragonal in cross section compressed pos-
teriorly, with a subcaudal groove present; anal folds present on sides
of cloaca; small glandular area behind insertion of hind limb, dark
in color.

Parvimolge richardi sp. nov.

Type: R. C. T.* no. 1436 collected at Isla Bonita (American Cin-
chona Plantation), Atlantic slope of Volcan Poas at an elevation of
about 6,500 feet; Aug. 1, 1947, by Richard Clark Taylor.

Diagnosis: A member of the genus Parvimolge, but differing from
Parvimolge towTisendi in the absence of the series of enlarged dorsal
glands, but agreeing in the ossification of the skull and the modifi-
cation of the digits, diminutive body size, moderately enlarged nos-

Field number of Richard C. Taylor.

Taylor: Salamanders from Costa Rica 285

tril, presence of maxillary teeth, and absence of an orbito-labial
groove.

Snout rather pointed; digits grown together, tips of fingers and
toes pointed; tail not constricted at base; chin with an arched
groove; nostrils enlarged; thirteen costal grooves; five to six costal
folds between adpressed limbs; paravomerine teeth in a single
patch, rrot notched behind; vomerine teeth on strongly elevated
ridges ; twelve maxillary teeth on each side ; six premaxillary teeth.

Description of type: Female; snout bluntly pointed; eye very
large, its horizontal diameter a little more than one and two fifths
times length of snout; nostril circular, moderately enlarged; inter-
orbital width about equal to an eyelid; surface of head smooth, the
openings of skin glands scarcely evident save in a curved series on
frrehead, between and partly anterior to eyelids.

Gular fold with an irregular groove passing up on each side of
head and meeting its fellow on the median line; on chin a groove
arching forward (apparently without a groove at base) ; on each
side of the base, a groove passes up across the angle of the jaw to
the dorsal surface of the head, then turns diagonally backward and
joins its fellow on the mid line somewhat in front of the nuchal
groove; a dim groove runs from eye back to the nuchal groove on
side of head; thirteen costal grooves; fold caused by the posterior
extension of the hyoid cartilage reaching as far as the third costal
fold; paravomerine teeth form a single elongate patch, widened and
rounded posteriorly, coming nearly to a point anteriorly, separated
from vomerine series by a short hiatus; vomerine teeth on two ele-
vated ridges about eight on a side, narrowly separated mesially;
maxillary teeth about 12-12 not reaching back as far as the middle
of eye; six premaxillary teeth; about 16-16 mandibular teeth.

Adpressed limbs separated by six costal folds. Limbs small, weak,
the hands and feet small; digits grown together rather than
"webbed." Third finger proportionately very large, the pointed
tip extending more than a third of its length beyond other fingers;
tip of second finger free, while those of the first and fourth are not or
scarcely free. Foot with the first and fifth toes not free; first joint
of the second and fourth with terminal joint free; third toe propor-
tionally large, the tip extending beyond other toes for nearly one
third of its length.

Sides of the cloaca with smooth diagonal folds.

Color: Above brownish lavender, darker on the head, especially
on upper eyelids, dorsally growing lighter to the base of the tail ; a

286 The University Science Bulletin

very indefinite darker lateral line; lower part of sides as light or
lighter than dorsum ; under side of chin and abdomen light purplish
with very numerous white or cream flecks; anal region light; tail
dark with numerous lighter flecks or vermiculations. The small
glandular spot behind insertion of leg scarcely discernible; white
flecks below nostril and eye; a white line on tip of snout.

Measurements: Total length, 49.5; snout to vent, 28; snout to
foreleg, 7.6; tail, 22; head width, 4.9; length of head to gular fold
(ventral), 5; length to gular fold (lateral), 5.85.

Remarks: This specimen was taken near the base of a stump un-
der bark in company with specimens of another undescribed species.

The absence of tha enlarged glands on the back show that, while
generically related, pcmjvnwtge and richardi are widely distinct spe-
cifically. The bluntly pointed snout likewise is a differentiating
character easily discerned.

The species is named for Richard Clark Taylor, its discoverer.

Oedipina serpens sp. nov.

Type: University of Kansas Museum of Natural History no.
23815 ^ ; collected at Morehead Finca, 5 miles southwest of Tur-
rialba, Costa Rica, July 21, 1947, by Edward H. Taylor.

Diagnosis: A very large species of the genus; snout to end of
vent, 74.5 mm.; tail more than twice head-body length; width of
head in head body length, 9.08 times; head length to gular fold
(median), 6.1 times; snout rather elongate, truncate not "sharp";
eye small; snout strongly overhanging lower jaw; digits grown to-
gether, free at tip, and rounded ; 8-9 costal folds between adpressed
limbs.

Description of the type: Snout rather elongate, truncate, the dor-
sal surface curving without trace of canthus rostralis; median part
of snout and interorbital region most elevated; eye very small, its
diameter contained in snout length twice, or very slightly less; a
small swelling below nostril on lip; posterior part of eyelids not
tucked under a diagonal fold; nostrils very small, lunate in shape;
width of upper eyelid in smallest interorbital distance, three times.

Gular fold strong, curving forward mesially ; from sides of fold a
nuchal groove runs up on side of neck but fails to meet its fellow on
the middorsal line ; a distinct postorbital groove runs back from the
eye directed slightly downward, then straight back to beginning of
nuchal groove; a groove crosses chin in front of the gular fold,
passes behind angle of mouth and reaches to dorsal surface of occi-

Taylor: Salamanders from Costa Rica 287

put; a short groove about midway between this and the nuchal
groove laterally terminating at postorbital groove; only a faint sug-
gestion of an arched groove on chin; no orbitolabial groove; no
median dorsal groove; costal grooves nineteen, traceable to midven-
tral line, and to near middle of back; presumably a constriction
at base of tail (tail severed) ; skin very smooth; limbs small, sep-
arated by nine costal folds when adpressed; arm adpressed reaches
about two thirds distance to eye; fingers broad, grown together
rather than "webbed," tips free with indistinct pads below; first
finger without free tip, rather well developed; median finger rounded
at tip; the other two fingers slightly narrowed at tip; toes very
broad, grown together, the tips rounded, terminal joint free on all
sav„ inner, with small pads below tips'; a white glandular spot be-
hind leg insertion; walls of cloaca with very numerous papillae.
Tail (severed and terminal portion missing) thick at base, tapering
gradually ; 52 -|- caudal grooves present.

Paravomerine teeth in two series narrowly separated mesially;
vomerine teeth in two series, 15-16, which originate much outside
outer level of choanae, run straight across mouth for greater part of
their length, then curve back, separated from each other by a dis-
tance equal to space between four teeth* and separated from the
paravomerine series by only a slightly greater distance. Choanae
small, their length contained in distance between them about five
times; three premaxillary teeth piercing gums; maxillary teeth
r.bout 50-50 ; mandibular teeth about 55-55 ; a strong free sublingual
fold.

Posterior extension of hyoid reaches to level of arm insertion.

Color: Dorsal and ventral color, grayish slate (under a lens the
circular glands are whitish gray, the intervening space black) ; spots
behind leg insertion bluish white ; under surface of limbs somewhat
brownish with some small whitish flecks; tip of snout with some
indefinite lighter flecks; subnarial swellings and a minute line on
edge of upper lip- cream; lower eyelid and part of edge of upper
whitish ; medial area on chin brownish with cream flecks ; hands and
feet grayish on palms and soles.

Measurements {in mm.): Snout to posterior end of vent, 74.5;
snout to arm insertion, 20; axilla to groin, 47; arm, 10; leg, 13.2;
head width, 8.8; head length to gular fold (medial), 12; head length
to gular fold (lateral), 15; width of body, 9; width of tail base, 7;
length of tail, 136 (missing part estimated at about 50 mm.).

Remarks: The specimen was found under a log in rather soft mud

288 The University Science Bulletin

at the edge of a stream bank. It was especially active and elusive
and escaped into a pile of large chips from which it was recaptured
with considerable effort. The tail was broken and part lost, perhaps
by my stepping on chips under which it was moving. Search was
made for the lost portion of the tail but it was not found.

Relationship: The relationship apparently is with Oedipina col-
laris Stejneger. The following table of contrasting characters show
the major differences between collaris and the one here described.

Oedipina collaris Oedipina serpens

1. Snout sharp. 1. Snout blunt, strongly truncate.

2. Head width in head body length, 2. Head width in head body length,
7.5 times. 8.4 times.

3. Head length in head body length, 3. Head length in head body length,
4.5 times. 4.9 — 6.2 times.*

4. Skin granular. 4. Skin smooth.

5. Vomerine teeth about nine. 5. Vomerine teeth fifteen.

6. Vomerine teeth extend straight 6. Vomerine teeth straight for part
across mouth. of series then curving back.

7. Fingers and toes short, scarcely 7. Fingers and toes long, strongly
flattened. flattened.

8 Paravomerine (parasphenoid) 8. Paravomerine teeth separated

teeth separated from vomerine from vomerine teeth by one-third

teeth by half length of latter length of one of latter series,

series. 9. Nine costal folds between ad-

9. Eleven costal folds between ad- pressed limbs.

pressed limbs. 10. Last joints of second, third and

10. Last joint of the third and fourth fourth free of membrane.
toes free of membrane. 11. These markings not present.

11. Large dark preocular spot and a
narrow black postocular streak.

* The gular fold curves strongly forward on middle of throat; head measured to this point,
its length is 12 mm., to the side of head the measurement is 15 mm.

LITERATURE CITED

1876. Cope, Edward D. On the Batrachia and Reptilia of Costa Rica. Joum.

Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876, pp. 93-157, pis. 23-28.
1887. Cope, Edward D. Catalogue of Batrachians and Reptiles of Central

America and Mexico. U. S. Nat. Mus. Bull. No. 32, 1887, pp. 1-96.
1926. Dunn, Emmet Reid. The Salamander of the Family Plethodontidae.

Smith College, 1926, Northampton, Mass., pp. 1-441, figs. 1-86.
1885-1902. GiJNTHER, Albert C. L. G. Biologia Centrali-Americana ; Reptilia

and Batrachia.

1933. Schmidt, Karl P. New Reptiles and Amphibians from Honduras. Zool.

Series Field Mus. Nat. Hist., vol. 20, 1933, pp. 15-22.
1936. Schmidt, Karl P. Guatemalan Salamanders of the genus Oedipus.
Zool. Ser. Field Mus. Nat. Hist., vol. 20, 1936, pp. 135-166, figs. 14-19.

1934. Wbtstein, Otto. Ergebnisse der osterreichischen biologischen Costa

Rica Expedition 1930. Die Amphibien und Reptilien. Sitz. Akad.
Wiss. Wien; Math-nat. Klasse., Abt. 1, Bd. 143, Heft 1-2, 1934, pp.
1-39.

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University of Kansas Science Bulletin - Vol. XXXIII - Part II

March 20, 1950
Lawrence, Kansas

ANNOUNCEMENT

The University of Kansas Science Bulletin (continuation of the
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The actual date of publication (i. e., mailing date) of many of the
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markedly from the dates bourne on the covers of the publication or
on the covers of the separata that it seems wise to offer a corrected
list showing the mailing date. The editor has been unable to verify
mailing dates earlier than 1932. Separata were issued at the same
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Vol. XXXII— Nov. 25, 1948.
Vol. XXXIII, Pt. I— April 20, 1949.

Editor Edward H. Taylor

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Contents of Volume XXXIII, Part II

No. PAGE)

7. Studies on Kansas Mosses I R. L. McGregor, 291

8. Type Localities of Mexican Reptiles and Amphibians

Hobart M. Smith and Edward H. Taylor, 313

9. Anomalies Found in a Series of Fetal and Newborn

Puppies Homer B. Latimer, 381

10. The Bees of the Genus Proteriades (Hymenoptera,

Megachilidae) . . . .P. H. Timberlake and C. D. Michener, 387

11. Second Contribution to the Herpetology of

San Luis Potosi Edward H. Taylor, 441

12. On the Status of the Family Desmognathidae

(Amphibia, Caudata) E. Ivan Soler, 459

13. Ceylonese Lizards of the Family Scincidae

Edward H. Taylor, 481

14. A Brief Review of Ceylonese Snakes. . . .Edward H. Taylor, 519

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Vol. XXXIII, Pt. II] March 20, 1950

[No. 7

Studies on Kansas Mosses I

BY

R. L. McGregor,
University of Kansas, Lawrence

Abstract: Of the total of 167 species and varieties of mosses, which have
been reported in the literature from Kansas, 73 are considered doubtful and
indeterminable. This leaves 94 collected by the author or represented by
specimens in various herbaria. The present paper reports 51 mosses not
previously listed for Kansas, giving a total of 145 species and varieties for
which collected specimens exist. These species are listed together with their
common habitats and the counties from which they have been collected.
Much field work and taxonomic study is necessary before our knowledge of
the Kansas moss flora becomes complete.

The study of Kansas bryophytes has been almost totally neglected
in the past. Except for a brief period of activity between 1884 and
1894 little has been accomplished. During those years several in-
dividuals collected and recorded in the literature a considerable
number of mosses. Their publications indicate the difficulties they
encountered in identifying and naming these plants. As a result,
certain of their reported species cannot now be recognized and only
a few of their specimens can be found. Errors made by these writers
in stating the authority for species increases the doubt as to just
what species they may have had.

The most recent paper on Kansas mosses is that of Gier (1949).
This paper gives a brief review of Kansas bryological work and re-
ports a few mosses previously unrecorded for the state, but for the
most part Gier's list is a compilation from literature. He has re-
vised the nomenclature and has brought it into agreement with that
employed by Grout (1928-1940). He brings together previously
published lists, but he has not added materially to our knowledge
of Kansas mosses.

(291)

292 The University Science Bulletin

Attempts at locating specimens from early collections of Kansas
mosses have proved almost futile. A small number of specimens
were found in the herbarium at Kansas State College, Manhattan,
and a few were obtained from Prof. Agrelius of Emporia State
Teachers College, Emporia, Kansas. Gier (1949) found some speci-
mens in the Chicago Museum of Natural History and lists a few
other herbaria that contain specimens from Kansas. Grout (1928-
1940) reported a few specimens, mostly as being in the herbarium
of the New York Botanical Garden. It is undoubtedly true that
other specimens exist, but such material is unknown to the author.

At the beginning of this work it was apparent that an adequate
treatment of the Kansas moss flora would require a fresh start at
collecting and naming. With this in mind the author has collected
in 82 of the 105 counties in the state. Such collections have, how-
ever, been made in conjunction with other field work and have not
been made with mosses as the primary item. Thus the several hun-
dred moss collections which form the basis for this paper do not
represent a careful survey of the state.

One reason for the neglect of Kansas bryology may have been
that the climate of the area would not be expected to favor an ex-
tensive moss flora. We find that Renauld and Cardot (1892) re-
ferred to Kansas as ". . . an area being in a region of low rain-
fall, high temperature, nearly treeless, and that frequent, sudden,
often severe weather changes occur." This, while only partly true,
is still the general opinion in regard to the state. However, the real
reason lies in the fact that the systematic botanists of the state
have been fully occupied with higher plants, a study that is yet far
from being finished.

It is not difficult to find mosses in the state. The eastern half
of the state has many limestone and sandstone outcroppings that
are usually wooded. In such places one can quickly fill a collecting
bag with specimens. The few remaining prairies, particularly the
rocky ones in hilly country, yield specimens. Creek banks and other
waste areas have their own moss flora. In the western half of the
state, rocky hillsides and rock outcroppings yield fewer species, but
those present are plentiful. In the same area one finds specimens
growing on the open plains, on roadside banks, around springs and
ponds and along partially wooded or unwooded stream banks.

In the present paper only those mosses for which known speci-
mens exist will be reported as occurring in the state. All other pub-
lished records are considered doubtful, since it is impossible to

McGregor: Studies on Kansas Mosses I 293

know whether the plants were correctly named. It is hoped that
anyone knowing of unreported species or possessing specimens will
bring them to the author's attention. It is also hoped that other
workers in the state will collect mosses and either report their find-
ings or send specimens in for determination. Much remains to be
done with the Kansas moss flora including the listing of species, their
distributions, and with certain species, taxonomic problems that are
beyond the scope of this paper.

During the progress of the present work I have drawn freely upon
others for assistance. Without their help this study would have
been impossible. In this regard I wish to thank Prof. Henry S.
Conard of Grinnell, Iowa, for making many determinations and of-
fering information whenever called upon. My thanks also are of-
fered to A. Leroy Andrews, Ithaca, N. Y. ; F. C. Gates, Kansas State
College, Manhattan ; F. U. G. Agrelius, Kansas State Teachers Col-
lege, Emporia; T. M. Sperry, Kansas State Teachers College, Pitts-
burg; S. V. Eraser, Aurora, Kansas; W. W. Holland, Walnut,
Kansas; W. H. Horr, R. H. Thompson and A, J. Mix of the Uni-
versity of Kansas; and to my wife, Dorothy McGregor, for her
valued assistance in collecting and preparing specimens for study
and preservation.

In the following list of species definitely known to occur in the
state the entries are arranged alphabetically by genera and species
following the nomenclature of Grout (1940). With each species is
given the habitat and information as to distribution. Unless other-
wise indicated all specimens are to be found in the herbarium at the
University of Kansas. Those specimens to be found in the herbarium
at Kansas State College, Manhattan, Kansas, are designated by
(KSC). Those at Kansas State Teachers College, Pittsburg, Kan-
sas, by (KSTCP). The rest are designated fully under the entry.
Amhlystegiella conjervoides (Brid.) Loeske

On moist limestone rocks. Cherokee Co., McGregor 1942.
Amblystegium americanum Grout

On moist soil near spring on rocky, prairie hillside. Scott Co., McGregor
2159, 2162.

Amblystegium juratzkanum Schimp.

On bark of elm and oak trees, fallen logs. Douglas Co., McGregor 2000;
Washington Co., McGregor 2044; Wyandotte Co., M. Reed April 22, 1891
(KSC).

Amblystegium, serpens (Hedw.) Bry. Etir.

On rocks and rotten logs. Ellsworth Co., McGregor 1497; Johnson Co.,
McGregor 2339.

294 The University Science Bulletin

Amblystegium varium (Hedw.) Lindb.

On rocks, soil, logs, bark of trees. Atchison Co., McGregor 1799; Brown Co.,
McGregor 1050; Butler Co., McGregor 1244, 2249; Chase Co., McGregor 1814;
Cherokee Co., McGregor 1942, 2363; Crawford Co., L. J. Gier 721 (KSTCP),
McGregor 1763, 1751; Douglas Co., McGregor 2279, 2322; Johnson Co., Mc-
Gregor 2344, 2388; Marshall Co., McGregor 2027; Miami Co., McGregor 1005,
1039; Morris Co., McGregor 1847; Nemaha Co., McGregor 2007, 2010; Ottawa
Co., McGregor 1475; Riley Co., Kellerman May 27, 1887 (KSC) ; Washing-
ton Co., McGregor 2045; Wilson Co., McGregor 2376; Woodson Co., Mc-
Gregor 2457; Wyandotte Co., McGregor 2423.

Amblystegium varium, (Hedw.) Lindb. var. ovatum Grout

Moist soil below sandstone outcrop. Ottawa Co., McGregor 1322.

Amblystegium varium (Hedw.) Lindb. var. parvulum (Aust.) Grout
Soil at base of tree. Douglas Co., F. U. G. Agrelius, March 1906.

Anomodon atlenuatv^s (Hedw.) Huben.

Soil, rocks, bark of trees. Bourbon Co., McGregor 1776; Cherokee Co.,
McGregor 1637, 1641; Crawford Co., McGregor 2525; Johnson Co., McGregor
2332, 2333; Wyandotte Co., McGregor 2425.

Anomodon minor (Beauv.) Lindb.

Soil, rocks, bark of trees. Atchison Co., McGregor 1801 ; Bourbon Co., Mc-
Gregor 1776, 2517; Douglas Co., McGregor 2280, 2323, 2446; Jefferson Co., Mc-
Gregor 1409; Johnson Co., McGregor 2329, 2345; Linn Co., McGregor 1957;
Lyon Co., McGregor 1593; Riley Co., Kellerman May, 1884 (KSC); Wyan-
dotte Co., McGregor 2424.

Anomodon rostratv^s Schimp.

On limestone rocks. Bourbon Co., Bennett Nov. 14, 1892 (KSC), McGregor
1776; Johnson Co., McGregor 2329; Miami Co., McGregor 2477; Riley Co.,
Kellerman April 5, 1888 (KSC).

Aphanorhegma serratum (Hook, and Wils.) Sull.

Moist soil, creek banks. Cherokee Co., McGregor 2559; Decatur Co., Mc-
Gregor 2104; Douglas Co., McGregor 2463; Leavenworth Co., McGregor 2260;
Miami Co., McGregor 2471.

Aschisma kansanum Andrews

This species was described by Andrews (1915) from material collected by
E. Hall in western Kansas. The exact location is unknown except that it was
collected on the prairies of western Kansas. The type specimen is in the
herbarium of the New York Botanical Garden and the Sullivant and James
herbarium.

Astomum muhlenbergianum (Sw.) Grout

Soil, wooded hillside. Jackson Co., McGregor 1973

Atrichum angustatum (Brid.) Bry. Eur.

Sandy or rocky soil, roadside banks, woods and thickets. Allen Co., Mc-
Gregor 1783, 1785; Anderson Co., McGregor 1577, 2497; Atchison Co., Mc-
Gregor 1804; Chautauqua Co., McGregor 1196; Cherokee Co., McGregor 1020,
1646, 2354; Cloud Co., Fraser Sept. 13, 1948; Crawford Co., Gier 733 (KSTCP),

McGregor: Studies on Kansas Mosses I 295

McGregor 1768, 2527; Douglas Co., McGregor 1045, 1054; Franklin Co.,
Meeker Nov. 21, 1892, Herrick Nov., 1892 (both specimens in Chicago Natural
History Museum); Johnson Co., McGregor 1330; Labette Co., McGregor 1619;
Leavenworth Co., McGregor 2269; Miami Co., McGregor 1011; Montgomery-
Co., McGregor 2388; Ottawa Co., McGregor 1472; Pottawatomie Co., Gates
17288 (KSC), Kellerman June 22, 1888 (KSC), Reed July 7, 1893 (KSC) ;
Riley Co., Kellerman Nov. 5, 1888 (KSC) ; Saline Co., Henry 1886 (Chicago
Natural History Museum); Shawnee Co., McGregor 1229; Wilson Co., Mc-
Gregor 2380.

Atrichum macmillani (Holz.) Frye

On soil in woods. Crawford Co., W. W. Holland 97 (KSC) ; Miami Co.,
McGregor 2592.

Atrichum undulatum (Hedw.) Beauv.

Sandy soil in woods. Washington Co., McGregor 2054.

Atrichum undulatum (Hedw.) Beauv. var. minus (Lam. and De Cand.) Web.
and Mohr.
Sandy soil below sandstone outcrop. Moist. Washington Co., McGregor
2043.

Atrichum undulatum (Hedw.) Beauv. var. selwynii (Aust.) Fiye

This variety is included here on the authority of Grout (1928-1940). The
county in which it was found is unknown to me though Reed (1894) listed
it from Saline County.

Aulacomnium heterostichum (Hedw.) Bry. Eur.

Sandy soil. Oak woods. Woodson Co., McGregor 2456.

Barhula cruegeri Sond.

Limestone rocks, clay soil. Dickinson Co., McGregor 1834; Douglas Co.,
McGregor 2321; Nemaha Co., McGregor 2014; RepubHc Co., McGregor 2085.

Barbida fallax Hedw.

Sandy roadside bank. Sumner Co., McGregor 2239.

Barhula unguiculata Hedw.

To bo found in nearly any type of habitat. Allen Co., McGregor 2510;
Clark Co., McGregor 2208; Clay Co., McGregor 2072; Cloud Co., McGregor
2070; Crawford Co., Gier 724 (KSTCP) McGregor 2526; Decatur Co., Mc-
Gregor 2133; Dickinson Co., McGregor 1841; Finney Co., McGregor 2181;
Hamilton Co., McGregor 2189; Jewell Co., McGregor 2090; Kearny Co.,
McGregor 2187; Leavenworth Co., McGregor 2267; Marshall Co., McGregor
2038; Nemaha Co., McGregor 2015; Osage Co., McGregor 1574; PhilHps Co.,
McGregor 2100; Pratt Co., McGregor 2217; Rawhns Co., McGregor 2139;
Repubhc Co., McGregor 2086, 2087; Riley Co., Kellerman March 24, 1884
(KSC) ; Smith Co., McGregor 2093.

Bartramia pomijormis Hedw.

On soil in woods, thickets and along stream banks. Chautauqua Co., Mc-
Gregor 1193; Cherokee Co., McGregor 1638, 1952; Douglas Co., McGregor
1222; Franklin Co., Herrick Nov., 1892 (KSC); Greenwood Co., McGregor
1911; Jackson Co., McGregor 1976; Linn Co., McGregor 1952.

296 The University Science Bulletin

Brachythedum fleoncaule Ren. and Card.

Rocky, sandy soil. Woods and thickets. Cloud Co., Fraser Sept. 13, 1948;
Crawford Co., McGregor 1764.

Brachythedum oxydadon (Brid.) Jaeger and Sauerb.

Rocky, sandy soil. Woods and thickets. Anderson Co., McGregor 2497;
Bourbon Co., McGregor 2517; Cloud Co., McGregor 2071, Fraser Sept. 13,
1948; Crawford Co., McGregor 2527; Ellsworth Co., McGregor, 1496; Leav-
enworth Co., McGregor 1392; Miami Co., McGregor 1010; Nemaha Co., Mc-
Gregor 2004; Republic Co., McGregor 2082; Riley Co., Reed Aug. 12, 1891
(KSC); Sumner Co., McGregor 2237; Wilson Co., McGregor, 2372.

Brachythedum oxydadon (Brid.) Jaeger and Sauerb. var. dentaium (Lesq.
and James) Grout.
Sandy soil, rocks, base of trees, creek bank. Cherokee Co., McGregor
2351; Crawford Co., McGregor 2525; Doniphan Co., McGregor 1808; Marshall
Co., McGregor 2029; Washington Co., McGregor 2040, 2041.

Brachythedum, salebrosum (Web. and Mohr.) Bry. Em-.

Sandy soil, rocky wooded hillsides, stream banks. Crawford Co., Gier 728
(KSTCP); Labette Co., McGregor 1618; Mitchell Co., McGregor 1993; Wyan-
dotte Co., Reed May 13, 1891 (KSC).

Bruchia brevifolia Sull.

On exposed, dry, sandy soil. Wilson Co., McGregor 1384.

Bruchia sullivanti Aust.

On sandy soil in oak woods. Chautauqua Co., McGregor 1370; Crawford
Co., reported by Gier (1949), specimens in herbarium at William Jewell Col-
lege, Liberty, Missouri.

Bryum argenteum (L.) Hedw.

Occurs on a wide range of habitats from soil in swamps, to open places in
prairie and on rock outcrops. Anderson Co., Thompson Feb. 2, 1948; Chey-
enne Co., McGregor 2145; Crawford Co., reported by Gier (1949); Doniphan
Co., McGregor 1966; Douglas Co., McGregor 2454; Kearny Co., McGregor
2186, 2187; Kingman Co., McGregor 2227; Labette Co., McGregor 1608, 1609;
Leavenworth Co., McGregor 1387; Lincoln Co., McGregor 1985; Lyon Co.,
Agrelius May 30, 1912, McGregor 1589; Morris Co., McGregor 1849; Neosho
Co., Holland 94 (KSC); Ottawa Co., McGregor 1317; Pratt Co., McGregor
2218; Reno Co., Voth Aug. 22, 1944 (in Herbarium Chicago Museum Natural
History); Saline Co., McGregor 1485; Washington Co., McGregor 2060, 2062;
Wabaunsee Co., McGregor 1856; Wilson Co., reported by L. J. Gier (1949),
specimen in herbarium William Jewell College, Liberty, Missouri.

Bryum argenteum, (L.) Hedw. var. lanatum (Beauv.) Bry. Eur.

Rocky hillsides and open places in prairie. Anderson Co., Thompson Feb.
2, 1948; Clay Co., McGregor 2074; Jefferson Co., McGregor 1412; Kingman
Co., McGregor 2220.

Bryum bicolor Dicks.

On rocks, sandy banks, creek banks, open places in prairie. Chase Co.,
McGregor 1810; Cloud Co., McGregor 2064; Decatur Co., McGregor 2105;

McGregor: Studies on Kansas Mosses I 297

Dickinson Co., McGregor 1835; Harper Co., McGregor 2235; Pratt Co., Mc-
Gregor 2217; Rawlins Co., McGregor 2137, 2139; Reno Co., Voth Aug. 22,
1944, reported by Gier (1949); Republic Co., McGregor 2086; Wabaunsee
Co., McGregor 1855.

Bryum caespiticium (L.) Hedw.

On sandstone and limestone rocks, rocky soil, creek banks, open places in
prairie, sagebrush flats and roadside banks. Barber Co., W. H. Horr 3114;
Barton Co., McGregor 1304, 1305; Cherokee Co., McGregor 1026; Clark Co.,
McGregor 2210; Crawford Co., Holland 92 (KSC) ; Decatur Co., McGregor
2103, 2133; Douglas Co., McGregor 1034, 2454; Grant Co., W. H. Horr 3081;
Harper Co., McGregor 2235; Leavenworth Co., McGregor 1391; Lincoln Co.,
McGregor 1984; Lyon Co., McGregor 1590; Marion Co., McGregor 1817;
Meade Co., McGregor 2201; Montgomery Co., McGregor 1176; Nemaha Co.,
McGregor 2003; Phillips Co., McGregor 2099; Pratt Co., McGregor 2218;
Republic Co., McGregor 1326; Washington Co., McGregor 2050, 2060; Wyan-
dotte Co., Reed May 5, 1892 (KSC).

Bryum cuspidatum (Bry. Eur.) Schimp.

On limestone and sandstone rocks; on soils in woods, prairie, stream banks
and swampy areas. Butler Co., McGregor 2248; Dickinson Co., McGregor
1834, 1842; Douglas Co., McGregor 1041; Marshall Co., McGregor 2034;
Nemaha Co., McGregor 2006, 2009; Rawlins Co., McGregor 2140; Republic
Co., McGregor 2084, 2087; Scott Co., McGregor 2177; Washington Co., Mc-
Gregor 2049.

Bryum gem.miparum DeNot.

On moist soil and rocks below a spring. Scott Co., McGregor 2161.

Bryum pendulum (Hornsch.) Schimp.

Sandstone rocks and sandy soil. Ellsworth Co., McGregor 1505, 1508;
Saline Co., McGregor 1483, 1494.

Bryum pseudotriquetrum (Hedw.) Schwaegr.

On logs, sandstone and limestone rocks, soil in woods. Cherokee Co., Mc-
Gregor 2362; Crawford Co., 1765, 1766; Miami Co., McGregor 1010; Nemaha
Co., McGregor 2008; Wilson Co., McGregor 2373.

Campylium chrysophyllu?n (Brid.) Bryhn.

On soil in woods, below rock outcrops, roadside banks and occasionally at
base of grasses in prairie. Atchison Co., McGregor 1802, 1804; Cherokee Co.,
McGregor 1020, 1025, 1644; Clay Co., McGregor 2075; Cloud Co., McGregor
2068; Crawford Co., Gier 727 (KSTCTP) ; Doniphan Co., McGregor 1808;
Douglas Co., McGregor 2301; Leavenworth Co., McGregor 1436, 1437; Miami
Co., McGregor 1008, 1011; Republic Co., McGregor 2078, 2081; Riley Co.,
Kellerman May 27, 1887 (KSC); Washington Co., McGregor 2042.

Campylium chrysophyllum (Brid.) Bryhn var. brevifoUum (Ren. and Card.)
Grout.

On rocky or sandy soil. Cherokee Co., McGregor 1764; Ellsworth Co.,
McGregor 1496, 1498; Leavenworth Co., McGregor 1395; Montgomery Co.,
McGregor 1178; Ottawa Co., McGregor 1469; Saline Co., McGregor 1491.

298 The University Science Bulletin

Campylium hispidulum (Brid.) Mitt.

Tree bark, sandy soil, stream banks. Cherokee Co., McGregor 1021, 1023,
1027, 1643; Crawford Co., Gier 723 (KSTCP), McGregor 1752, 1756, 1763;
Douglas Co., McGregor 1040; Leavenworth Co., McGregor 1398, 1399; Re-
public Co., McGregor 2081a; Riley Co., Reed June 30, 1892 (KSC).

Ceratodon purpureus (Hedw.) Brid.

On sandstone rocks, sandy soil, rocky soil in woods, roadside banks and
around base of grasses in prairie. Brown Co., McGregor 1049; Clark Co.,
McGregor 2209; Cloud Co., McGregor 2064; Cowley Co., W. H. Horr 3168;
Crawford Co., McGregor 1767; Douglas Co., Agrelius May 1, 1906; Ellsworth
Co., McGregor 1510; Leavenworth Co., McGregor 1391, 1394; Lyon Co., Mc-
Gregor 1590; Nemaha Co., McGregor 2005; Ottawa Co., McGregor 1317;
Rawlins Co., McGregor 2138; Republic Co., McGregor 1326, 1327; Riley Co.,
Reed Aug. 12, 1891 (KSC); Saline Co., McGregor 1481; Shawnee Co., Mc-
Gregor 1228; Wyandotte Co., Reed May 5, 1891 (KSC).

Chamberlainia acuminata (Hedw.) Grout.

On limestone rocks, tree bark, and soil at base of trees. Anderson Co.,
McGregor 2493; Douglas Co., McGregor 2274; Johnson Co., McGregor 2343;
Marshall Co., McGregor 2028; Mitchell Co., McGregor 1998; Morris Co.,
McGregor 1846; Pottawatomie Co., Reed Nov. 6, 1892 (KSC); Reno Co.,
McGregor 2225; Republic Co., McGregor 2083.

Climacium americanum Brid.

Sandstone rocks and soil in oak woods. Elk Co., McGregor 1923; Green-
wood Co., McGregor 1902.

Cratoneuron filicmum (Hedw.) Roth.

On soil among rushes and sedges in marsh. Atchison Co., McGregor 1017.

Desmotodon ohtusifolius (Schwaegr.) Jur.

Predominantly on sandstone rock, rarely on limestone. Occasionally on
sandy alluvial soil. Allen Co., McGregor 1788; Barton Co., McGregor 1304,
1305; Chase Co., McGregor 1812; Cheyenne Co., McGregor 2145; Crawford
Co., Holland 90 (Collector's herbarium); Douglas Co., McGregor 1043; Ells-
worth Co., McGregor 1511; Leavenworth Co., McGregor 1387, 2260; McPher-
son Co., McGregor 1820; Neosho Co., McGregor 1256; Ottawa Co., Mc-
Gregor 1317; Rawlins Co., McGregor 2135; Rice Co., McGregor 1831; Saline
Co., McGregor 1487; Washington Co., McGregor 2058, 2061; Wilson Co.,
McGregor 2381.

Desmatodon plinthobius Sull. and Lesq.

On sandstone and limestone rocks and soil near such rocks. Butler Co.,
McGregor 2244a; Chase Co., McGregor 1813; Dickinson Co., McGregor 1839;
Harper Co., McGregor 2232; Ottawa Co., McGregor 1318.

Desmatodon porteri James.

On limestone rocks. Atchison Co., McGregor 1796.

Dicranella heteromalla (Hedw.) Schimp.

On soil in woods. Occasionally on rocks. Allen Co., McGregor 1784;
Crawford Co., McGregor 1761; Douglas Co., McGregor 2302; Elk Co., Mc-

McGregor: Studies on Kansas Mosses I 299

Gregor 1921; Greenwood Co., McGregor 1904; Leavenworth Co., McGregor
1390, 2266; Montgomery Co., McGregor 2391.

Dicranella varia (Hedw.) Schimp.

On rocks, sandy alluvial soil. Allen Co., McGregor 2508; Anderson Co.,
McGregor 2490; Crawford Co., McGregor 2528; Douglas Co., McGregor 2306;
Leavenworth Co., McGregor 2263; Washington Co., McGregor 2052.

Dicranum condensatum Hedw.

Sandy soil in oak woods. Cliautauqua Co., McGregor 1201; Elk Co., Mc-
Gregor 1918; Montgomeiy Co., McGregor 1173.

Dicranum scoparium Hedw.

Moist soil in woods. Chautauqua Co., McGregor 1201; Cherokee Co., Mc-
Gregor 1018, 1944; Doniphan Co., McGregor 1971; Jackson Co., McGregor
1980; Leavenworth Co., McGregor 1390, 1432, 2259, 2261; Linn Co., Mc-
Gregor 1953; Montgomery Co., McGregor 1173.

Didymodon rigidulus Hedw.

On rocky soil. Logan Co., McGregor 2156.

Didymodon tophaceus (Brid.) Jur.

On soil near a spring in prairie. Scott Co., McGregor 2158, 2160.

Didymodon trifarius (Hedw.) Brid.

This species is incuded here on the authority of Gier (1949) and a personal
letter from Prof. Gier. He found a specimen of this species in the Chicago
Natural History Museum collected by W. R. Maxon in April, 1912. The locality
was not given on the label except for the name of the state.

Ditrichum pallidum (Hedw.) Hampe.

Moist soil in woods. Occasionally found among rocks in prairies of central
Kansas. Anderson Co., McGregor 1578; Chautauqua Co., McGregor 1190;
Cherokee Co., McGregor 1029, 1031; Crawford Co., Holland 91 (KSC) ;
Douglas Co., McGregor 1045; Franklin Co., McGregor 1341; Labette Co.,
McGregor 1606; Leavenworth Co., McGregor 1437; Lincoln Co., McGregor
1982; Miami Co., McGregor 1008; Ottawa Co., McGregor 1324; Wilson Co.,
McGregor 1385, 2380, McClung May 9, 1891 (KSC) ; Woodson Co., McGregor
1253.

Drepanocladus aduncu^ (Hedw.) Warnst. var. kneiffi (Bry. Eur.) Warnst.
On soil. Crawford Co., Gier 725 (KSTCP).

Drepanocladus aduncus (Hedw.) Warnst. var. polycarpus (Bland.) Warnst.

Very moist sandy soil. Mitchell Co., McGregor 1997; Trego Co., Reed July
8, 1893 (KSC).

Drummondia prorepens (Hedw.) Jennings.

On tree bark and limestone rocks. Cherokee Co., McGregor 1649, 1931.

Entodon cladorrhizans (Hedw.) C. Muell.

On limestone rocks. Johnson Co., McGregor 2332.

Entodon compressus (Hedw.) C. Muell.

On logs, bark of trees and moist soil. Bourbon Co., McGregor 2518; Butler

300 The University Science Bulletin

Co., McGregor 2250; Douglas Co., McGregor 2309, 2318; Johnson Co., Mc-
Gregor 2335, 2337; Marshall Co., McGregor 2025; Morris Co., McGregor 1845;
Nemaha Co., McGregor 2021; Ottawa Co., McGregor 1475; Washington Co.,
McGregor 2053.

Entodon seductrix (Hedw.) C. Muell.

On limestone and sandstone rocks, fallen logs, base of trees and moist soil
in woods. Allen Co., McGregor 1786, 2505; Anderson Co., McGregor 1964,
2492; Bourbon Co., McGregor 1769; Chautauqua Co., McGregor 1199; Cher-
okee Co., McGregor 1941; Crawford Co., Holland 95 (KSC), Gier 718
(KSTCP), McGregor 1762; Douglas Co., Agrelius Nov. 3, 1906 and April
28, 1906, Thompson March 14, 1948, McGregor 1055, 2278, 2281; Jefferson
Co., McGregor 1411; Labette Co., McGregor 1621; Leavenworth Co., Mc-
Gregor 1393, 2258, 2265; Lincoln Co., McGregor 1986; Linn Co., McGregor 1955;
Miami Co., McGregor 1053, 2476; Saline Co., McGregor 1490; Wilson Co.,
McGregor 2375.

Eucladium verticillatum (Brid.) Br3^ Eur.

Very moist limestone rocks. Cherokee Co., McGregor 1625.

Eurhynchium hians (Hedw.) Jaeger and Sauerb.

On rocks, logs, soil and bark of trees. Cherokee Co., McGregor 1025, 1632;
Dickinson Co., McGregor 1840; Douglas Co., McGregor 2317; Johnson Co.,
McGregor 2336; Leavenworth Co., McGregor 1395, 1398; Miami Co., Mc-
Gregor 1013; Nemaha Co., McGregor 2012; Shawnee Co., L. D. Voile Oct. 1,
1948; Wabaunsee Co., McGregor 1854; Wyandotte Co., Reed May 1, 1891
(KSC).
Eurhynchium serrulatum (Hedw.) Kindb.

Moist soil in woods. On logs and bark of trees. Cherokee Co., McGregor
1021; Crawford Co., Gier 722 (KSTCP), McGregor 1755, 1758; Douglas Co.,
Agrelius March, 1906, McGregor 1048, 2277, 2304; Johnson Co., McGregor 2343;
Wyandotte Co., McGregor 2422, 2424.

Fabronia wrightii Sull.

Bark of elm trees. Crawford Co., McGregor 1759; Dickinson Co., Mc-
Gregor 1843; Douglas Co., McGregor 2281; Harvey Co., McGregor 2253;
Marshall Co., McGregor 2022.

Fissidens bu^hii Card, and Ther.

On soil in rocky wooded hillside. Anderson Co., McGregor 2495.

Fissidens cristatus Wils.

Moist limestone rocks. Cherokee Co., McGregor 1635.

Fissidens cristatus Wils. var. winonensis (Ren. and Card.) Grout.
Moist sandy soil in oak woods. Greenwood Co., McGregor 1903.

Fissidens julianvs (Mont.) Schimp.

Attached to rocks, sticks and other submerged objects in clear, slow-mov-
ing streams. Cherokee Co., McGregor 2361; Woodson Co., McGregor 2459.

Fissidens minutulus Sull.

Moist sandstone and limestone rocks. Also moist sandy banks. Atchison

McGregor: Studies on Kansas Mosses I 301

Co., McGregor 1799; Cherokee Co., McGregor 1943; Franklin Co., McGregor
1515; Leavenworth Co., McGregor 2264; Republic Co., McGregor 2080; Wil-
son Co., McGregor 2376, 2384.

Fissidens ohtusijolius Wils.

Moist sandstone and limestone rocks. Chase Co., McGregor 1812; Leaven-
worth Co., McGregor 1579, 2260, 2264; Washington Co., McGregor 2056, 2058.

Fissidens ohtusijolius Wils. var. kansanus Ren. and Card.

This variety was described by Renauld and Cardot (1890) from material
sent them by Dr. Joseph Henry of Salina, Kansas. This fact was mentioned
by Grout (1928-1940). I have not checked on the location of the type her-
baiium specimens. The type locality is Saline County.

Fissidens subbasilaris Hedw.

On moist limestone rocks. Cherokee Co., McGregor 1941 ; Douglas Co.,
McGregor 2307; Johnson Co.. McGregor 2340.

Fissidens taxifolius Hedw.

This species was reported from Crawford County Kansas State Park by
Gier (1940, 1949). I have seen a packet of his material on file in the her-
barium of Kansas State Teachers College, Pittsburg, Kansas.

Fontinalis missourica Card.

Gier (1949) and in personal correspondence with me says there is a packet
of this species in the herbarium of the Chicago Museum of Natural History.
The specimen was collected by E. Hall in 1870. According to Gier (1949) the
locality was given as "headwaters of the Neosho River." This would seem to
place the actual location of the collection to the northwest of Greenwood
County. Gier (1949) mentioned Greenwood County as a possible location.

Funaria calvescens Schwaegr.

Sandy soil. Wilson Co., McGregor 1276.

Funaria flavicans Mx.

Moist soil in woods. Chautauqua Co., McGregor 1191 ; Douglas Co., Mc-
Gregor 1033; Jefferson Co., McGregor 1414; Johnson Co., McGregor 1333;
Woodson Co., McGregor 1252.

Funaria hygrometrica Hedw.

Open places in woods especially recently burned over areas, roadside banks,
creek banks and abandoned fields. Also open places in prairie. Brown Co.,
McGregor 1049; Cherokee Co., McGregor 1022; Crawford Co., Gier 729
(KSTCP), Holland 99 (KSC), McGregor 1832; Douglas Co., McGregor 1044,
1421; Dickinson Co., McGregor 1837; Ellsworth Co., McGregor 1507; Finney
Co., McGregor 2182; Hamilton Co., McGregor 2189; Labette Co., McGregor
1609, 1610; Lyon Co., Agrelius June 29, 1912; McPherson Co., McGregor 1821;
Morris Co., McGregor 1849; Ottawa Co., McGregor 1323, 1324; Pottawatomie
Co., Reed July 7, 1893 (KSC); Rice Co., McGregor 1824, 1832; Riley Co.,
Norton 1893 (KSC); Saline Co., McGregor 1493; Washington Co., McGregor
2056; Woodson Co., McGregor 1793; Wyandotte Co., Reed April and May
1891 (KSC).

302 The University Science Bulletin

Grimmia alpicola Hedw.

On limestone rocks. Bourbon Co., McGregor 1770.

Grimmia apocarpa Hedw.

On dry exposed limestone and sandstone rocks. Anderson Co., Thompson
Feb. 2, 1948; Chautauqua Co., McGregor 1194; Clark Co., McGregor 2210;
Cowley Co., McGregor 2241; Lyon Co., McGregor 1594; Marshall Co., Mc-
Gregor 2026; Mitchell Co., McGregor 1996; Scott Co., McGregor 2176;
Wyandotte Co., Reed July 24, 1893 (KSC).

Grimmia laevigata (Brid.) Brid.

Dry exposed sandstone rocks. Cliautauqua Co., McGregor 1192, 1197;
Douglas Co., McGregor 2314; Saline Co., McGregor 1492; Wilson Co., Mc-
Gregor 2371.

Grimmia plagiopodia Hedw.

Dry rocks. Rice Co., McGregor 1825.

Grimmia raui Aust.

Found abundantly on a rock outcrop in western Kansas known commonly
as the mortar beds. Clark Co., McGregor 2207; Decatur Co., McGregor 2132;
Kearny Co., McGregor 2185; Logan Co., McGregor 2152, 2155; Meade Co.,
McGregor 2206; Rawlins Co., McGregor 2134; Scott Co., McGregor 2175.

Gymnostomum recurvirostrum Hedw.

On moist, rocky soil in woods. Cherokee Co., McGregor 2366.

Hedwigia ciliata Hedw.

On sandy soil and sandstone rocks. Allen Co., McGregor 1782; Chautauqua
Co., McGregor 1198; Doniphan Co., McGregor 1968; Douglas Co., McGregor
2315; Greenwood Co., McGregor 1910; Wilson Co., McGregor 2374, 2378.

Homo7nallium adnatum, (Hedw.) Broth.

Limestone and sandstone rocks. Tree bark. On soil in woods and roadside
banks. Allen Co., McGregor 1790; Bourbon Co., McGregor 2520; Cherokee
Co., McGregor 1019, 1636; Crawford Co., Holland 89 (KSC); Johnson Co.,
McGregor 2345, 2346; Labette Co., McGregor 1605, 1611; Lincoln Co., Mc-
Gregor 1983; Montgomery Co., McGregor 1174; Morris Co., McGregor 1851;
Wilson Co., McGregor 1383.

Hygroamblystegium irrigwxm (Wils.) Loeske.

On logs, base of trees, sandstone and limestone rocks, moist soil in woods.
Allen Co., McGregor 1791; Anderson Co., McGregor 1962; Cherokee Co., Mc-
Gregor 1030, 1926, 1929; Doniphan Co., McGregor 1970; Douglas Co., Agre-
lius Feb. 22, 1906, McGregor 1036, 1422, 1630, 2276; Ellsworth Co., McGregor
1509; Leavenworth Co., McGregor 1392, 1434; Miami Co., McGregor 1012,
1929; Rice Co., McGregor 1829; Riley Co., Swingle May 1890 (KSC), Keller-
man May 6, 1887 (KSC); Saline Co., McGregor 1495; Scott Co., McGregor
2163, 2164; Wilson Co., McGregor 2384 ; Woodson Co., McGregor 1250; Wyan-
dotte Co., McGregor 2429.

Hygroamblystegium orthocladon (Beauv.) Grout.

On sandstone rock and calcareous soil. Leavenworth Co., McGregor 1386;
Morris Co., McGregor 1848.

McGregor: Studies on Kansas Mosses I 303

Hyjmum curvijolium Hedw.

On soil, wooded hillside. Jackson Co., McGregor 1979.

Hypnum patientiae Lindb.

Limestone rocks and soil in woods. Cherokee Co., McGregor 1626, 2263,
2364; Jackson Co., McGregor 1978.

Leptobryum pyrijorme (Hedw.) S'chimp.

On sandstone and limestone rocks. McPherson Co., McGregor 1821; Ne-
maha Co., McGregor 20O2; Pottawatomie Co., Reed July 7, 1893 (KSC).

Leptodictyum riparium (Hedw.) Warnst.

Rotten logs, moist soil in woods, base of cattails and sedges in swamps.
Atchison Co., McGregor 1016; Brown Co., McGregor 1051; Cherokee Co.,
McGregor 1631; Crawford Co., Gier 726 (KSTCP) ; Douglas Co., McGregor
2308; Ellsworth Co., McGregor 1310; Jefferson Co., McGregor 1417; Meade
Co., McGregor 1302; Miami Co., McGregor 1039, 2479.

Leptodictyum trichopodium (Schultz) Warnst.

Moist soil especially in seepy areas or swamps. Atchison Co., McGregor
1015; Douglas Co., McGregor 1034, 2278; Ellsworth Co., McGregor 1507;
Leavenworth Co., McGregor 1393; Meade Co., McGregor 1302, 2198, W. H.
Horr 3110; Ottawa Co., McGregor 1325, 1473; Rice Co., McGregor 1827;
Saline Co., McGregor 1482, 1489; Woodson Co., McGregor 2457.

Leptodictyum trichopodium, (Schultz) Warnst. var. kochii (Bry. Eur.) Broth.
On moist sandy soil. Dickinson Co., McGregor 1834; Douglas Co., Mc-
Gregor 1224.

Leskea gracilescens Hedw.

Bark of trees, rotten logs, rarely on soil in woods. Allen Co., McGregor
2509; Anderson Co.. McGregor 2493; Bourbon Co., McGregor 1771; Butler
Co., McGregor 2250; Cherokee Co., McGregor 1634, 1639; Cloud Co., Mc-
Gregor 2063, 2066; Coffey Co., McGregor 1599; Comanche Co., W. H. Horr
3116; Crawford Co., Gier 719 (KSTCP); Dickinson Co., McGregor 1833
Douglas Co., McGregor 2309; Franklin Co., McGregor 1575; Harvey Co.
McGregor 2252; Jefferson Co., McGregor 1410; Jewell Co., McGregor 2088
Johnson Co., McGregor 2331; Leavenworth Co., McGregor 1388; Linn Co.
McGregor 1958; Lyon Co., McGregor 1595; Marion Co., McGregor 1816
Marshall Co., McGregor 2024; McPherson Co., McGregor 1818; Miami Co.
McGregor 2465; Morris Co., McGregor 1850; Nemaha Co., McGregor 2010
Osage Co., McGregor 1571; Reno Co., McGregor 2224; Republic Co., Mo
Gregor 2077; Sumner Co., McGregor 2238; Washington Co., McGregor 2059
Wabaunsee Co., McGregor 1852; Wyandotte Co., Reed April 2, 1891 (KSC),
McGregor 2424.

Leskea obscura Hedw.

On bark of elms, oaks and logs. Allen Co., McGregor 2509; Atchison Co.,
McGregor 1799; Coffey Co., McGregor 1600; Crawford Co., McGregor 1757;
Douglas Co., McGregor 1224, Agrelius, May 1, 1906; Geary Co., F. C. Gates
18890 (KSC); Labette Co., McGregor 1613; Lincoln Co., McGregor 1989;
Miami Co., McGregor 1006; Mitchell Co., McGregor 1990; Nemaha Co.,

304 The University Science Bulletin

McGregor 2005; Pottawatomie Co., Reed Nov. 6, 1891 (KSC), Reed Nov. 26,
1892 (KSC).

Leucobryum albidum (Brid.) Lindb.

On sandy soil, oak woods. Chautauqua Co., McGregor 1201.

Leucobryum glaucum. (Hedw.) Schimp.

On sandy soil, oak woods. Anderson Co., McGregor 2496; Cherokee Co.,
McGregor 1647; Crawford Co., McGregor 2524; Elk Co., McGregor 1922;
Greenwood Co., McGregor 1906; Jackson Co., McGregor 1974; Montgomery
Co., McGregor 2387.

Leucodon brachypus Brid.

Credited to Kansas by Grout (1928-1940). I do not know location of
specimen or locality from which collected.

Leucodon julaceous (Hedw.) Sull.

Moist limestone rocks. Anderson Co., McGregor 2498; Allen Co., Mc-
Gregor 2506; Cherokee Co., McGregor 1653.

Lindbergia brachyptera (Mitt.) Kindb. var. austinii (Sull.) Grout.

This variety is reported from Kansas by Grout (1928-1940). I do not
know locality from which collected or location of specimen. Reed (1894)
reported it from Saline County.

Mnium affine Bland.

Moist clay creek bank. Douglas Co., McGregor 2305.

Mnium cuspidatum Hedw.

On sandstone and limestone rocks, soil in woods, creek banks, fields, base
of trees and tree bark. Allen Co., McGregor 1787; Anderson Co., McGregor
1577; Atchison Co., McGregor 1804, 1797; Bourbon Co., McGregor 1773;
Chautauqua Co., McGregor 1200; Cherokee Co., McGregor 1028, 1650; Craw-
ford Co., Holland 87 (KSC), Gier 734 (KSTCP), McGregor 1752; Douglas
Co., McGregor 1047, Agrelius May 1, 1910, Thompson March 14, 1948; Ells-
worth Co., McGregor 1503; Jefferson Co., McGregor 1416, 1418; Johnson Co.,
McGregor 1332, 2344; Labette Co., McGregor 1617; Leavenworth Co., Mc-
Gregor 1396; Lincoln Co., McGregor 1981; McPherson Co., McGregor 1823;
Miami Co., McGregor 1013; Nemaha Co., McGregor 2011; Ottawa Co., Mc-
Gregor 1476; Rice Co., McGregor 1827; Riley Co., Kellerman May 25, 1887
(KSC), Jones May 18, 1892 (KSC); Saline Co., McGregor 1486; Shawnee
Co., McGregor 1227, L. D. Voile Oct. 1, 1948; Washington Co., McGregor
2039; Wabaunsee Co., McGregor 1853; Woodson Co., McGregor 1251; Wyan-
dotte Co., McGregor 2426.

Orthotrichum, diaphanum, Brid.

Bark of elm trees. Reno Co., McGregor 2223.

Orthotrichum, pumilum Dicks.

Bark of elm, hackberry and oak. Cherokee Co., McGregor 1021, 1627, 1634;
Cloud Co., McGregor 2069; Crawford Co., McGregor 1754, 1759; Dickinson
Co., McGregor 1844; Doniphan Co., McGregor 1967; Douglas Co., McGregor
2621; Marshall Co., McGregor 2023; Washington Co., McGregor 2059.

McGregor: Studies on Kansas Mosses I 305

Orthotrichum jmsillum Mitt.

Bark of elm and oak. Cherokee Co., McGregor 1627; Labette Co., Mc-
Gregor 1607; Reno Co., Voth Aug. 1944, specimen in herbarium at Chicago
Natural History Museum according to Gier (1949).

Orthotrichum sordidum Lesq. and James.

Tree bark. Crawford Co., Holland 93 (KSC).

Orthotrichum strangulatum Schwaegr.

On limestone and sandstone rocks. Cherokee Co., McGregor 1640; Mitchell
Co., McGregor 1994; Riley Co., Kellerman May 4, 1884 (KSC).

Phascum, cuspidatum Hedw. var. americanum Ren. and Card.

On sandy soil, oak woods. Shawnee Co., McGregor 1230.

Philonotis glaucescens (Hornsch.) Paris.

This species reported by Grout (1928-1940). I do not know location of
specimen or locality from which collected.

Philonotis gracillima Angstr.

This species was reported by Grout (1928-1940). I do not know location
of specimen or locality from which collected.

Philonotis marchica (Willd.) Brid.

Moist sandy soil in woods. Cherokee Co., McGregor 2365; Montgomery
Co., McGregor 2388.

Physcomitrium hookeri Hampe.

On moist soil, edge of swamp. Meade Co., McGregor 1313.

Physcomitrium kellermani E. G. Britton.

Sandy soil, creek banks and open places in prairie. Cowley Co., McGregor
2243; Douglas Co., McGregor 1223; Kingman Co., McGregor 2228; Norton
Co., McGregor 2101 ; Riley Co., Kellerman 1889 (this is the type specimen
on file in herb. N. Y. Botanical Garden) ; Rook Co., McGregor 2096.

Physcomitrium turbinatum (Mx.) Brid.

Moist sandy soil, creek banks, seepy or swampy areas, moist fields. Chau-
tauqua Co., McGregor' 1371 ; Crawford Co., Holland 88, 101 (KSC), Gier 731
(KSTCP); Douglas Co., McGregor 1035, 1042, 1423, Agrelius April 20, 1906;
Ellsworth Co., McGregor 1504, 1506; Franklin Co., McGregor 1340; Jefferson
Co., McGregor 1414; Johnson Co., McGregor 1331; Labette Co., McGregor
1615; Leavenworth Co., McGregor 1615; Lincoln Co., McGregor 1987; Mitchell
Co., McGregor 1991; Osage Co., McGregor, 1573; Ottawa Co., McGregor 1470;
Saline Co., McGregor 1484, 1489; Sedgwick Co., Carleton 131 (KSC); Wyan-
dotte Co., Reed May 5, 1891 (KSC), McGregor 1443.

Plagiothecium deplanatum (Sull.) Grout.

Moist soil, wooded hillside. Wyandotte Co., McGregor 2421.

Plagiothecium, geophilum^ (Aust.) Grout.

Bark of elm. Atchison Co., McGregor 1800; Miami Co., McGregor 1012;
Wyandotte Co., McGregor 2422, 2424.

19—90

306 The University Science Bulletin

Platygyiiwn repens (Brid.) Bry. Eur.

Sandstone rocks, logs, tree bark. Bourbon Co., McGregor 1772; Cherokee
Co., McGregor 1627, 2352; Jackson Co., McGregor 1977; Johnson Co., Mc-
Gregor 2341; Leavenworth Co., McGregor 2268; Mitchell Co., McGregor 1998;
Pottawatomie Co., Reed Nov. 6, 1892 (KSC).

Pleuridium subulatum (Hedw.) Lindb.

Sandy soil, woods, prairie. Johnson Co., McGregor 1334; Leavenworth Co.,
McGregor 1430; Washington Co., McGregor 2048.

Pleuridium sullivanti Aust.

Dry sandy soil in prairie. Woodson Co., McGregor 1249.

Pogonatum brachyphyllum (Rich.) Beauv.

Sandy soil in woods. Douglas Co., McGregor 1052; Neosho Co., Thompson
March 6, 1949; Wilson Co., McGregor 2383; Woodson Co., McGregor 2636.

Pohlia nutans (Hedw.) Lindb.

On sandstone rocks, sandy soil in woods. Allen Co., McGregor 1784;
Cherokee Co., McGregor 1646; Crawford Co., McGregor 1761; Douglas Co.,
Agrelius April 4, 1910; Leavenworth Co., McGregor 1390, 1435, 2261; Miami
Co., McGregor 1009; Pottawatomie Co., Reed July 7, 1893 (KSC); Wilson
Co., McGregor 2382.

Pohlia wahlenhergii (Webr. and Mohr.) Andrews.

Sandstone rocks, sandy soil, moist soil in woods. Douglas Co., McGregor
2273; Leavenworth Co., McGregor 2267; Mitchell Co., McGregor 1995; Ottawa
Co., McGregor 1474; Riley Co., Reed April 7, 1893 (KSC); Saline Co., Hancin
1320 (KSC), McGregor 1482; Washington Co., McGregor 2051; Woodson
Co., McGregor 2458.

Polytrichum commune Hedw.

Sandy soil in oak woods. Elk Co., McGregor 1919; Greenwood Co., Mc-
Gregor 1905, 1909.

Polytrichum juniperinum Hedw.

Rocky or sandy soil in woods. Chautauqua Co., McGregor 1189; Cherokee
Co., McGregor 1018, 1024; Doniphan Co., McGregor 1969; Elk Co., McGregor
1914; Leavenworth Co., McGregor 1431; Montgomery Co., McGregor 1172;
Pottawatomie Co., Reed July 7, 1893 (KSC).

Polytrichum- ohioense Ren. and Card.

Rocky or sandy soil in woods. Cherokee Co., McGregor 1651; Douglas Co.,
McGregor 1014.

Pterigoneurum subsessile (Brid.) Jur.

Open places in rocky prairie. Rawlins Co., McGregor 2141.
Pterigoneurum subsessile (Brid.) Jur. var. henrici (Rau) Wareham.

The type material, according to Grout (1928-1940), is in the herbarium of
the New York Botanical Garden and was collected by Joseph Henry in Saline
County.

McGregor: Studies on Kansas Mosses I 307

Ptychomitrium incurvum (Muhlenb.) Sull.

On sandstone rocks. Montgomery Co., McGregor 1174, 1179.

Pylaida selwynii Kindb.

Bark of elm trees. Anderson Co., McGregor 1963; Linn Co., McGregor
1951; Mitchell Co., McGregor 1992; Riley Co., Kellerman May 1884 (KSC).

Pyratnidula tetragona (Brid.) Brid.

Sandy soil in prairie. Dougla.s Co., McGregor Nov. 2, 1947.
Rhodohryum roseum (Bry. Eur.) Limpr.

On soil in woods. Jackson Co., McGregor 1975.
Sematophyllum carolinianum (C. Mull.) E. G. Britton.

On sandstone rocks. Montgomery Co., McGregor 2390.
Sphagnum capillaceum (Weiss) Schrank

Moist sandy soil on bank of small creek. Woodson Co., Thompson Sept.
1945, McGregor 2647.

Thelia asprella Sull.

Limestone rocks, moist soil, bark of elm. Cherokee Co., McGregor 1945;
Douglas Co., McGregor 2445; Greenwood Co., E. Hall (in herbarium at Chi-
cago Museum of Natural History according to a personal letter received from
L. J. Gier of Liberty, Missouri); Johnson Co., McGregor 2331; Linn Co., Mc-
Gregor 1956.

Thelia hirtella (Hedw.) Sull.

On limestone rocks. Allen Co., McGregor 2507; Anderson Co., McGregor
2494.

Thelia lescurii Sull.

On moist limestone rocks. Cherokee Co., McGregor 1652, 1654; Douglas
Co., Agrelius Nov. 3, 1906.

Thuidium delicatulum (Hedw.) Mitt.

Moist limestone rocks, sandy soil, moist soil in woods. Cherokee Co., Mc-
Gregor 1642; Elk Co., McGregor 1920; Greenwood Co., McGregor 1901.

Thuidiwm microphyllum (Hedw.) Best.

Bark of elm. Cherokee Co., McGregor 1021; Crawford Co., Gier 720
(KSTCP).

Thuidium recognitum (Hedw.) Lindb.

On limestone rocks. Anderson Co., McGregor 1961, 2491; Cherokee Co.,
McGregor 1638; Crawford Co., McGregor 2523; Linn Co., McGregor 1948.

Thuidium virginianum (Brid.) Lindb.

On logs and bark of trees. Crawford Co., McGregor 1751; Jefferson Co.,
McGregor 1515; Miami Co., McGregor 1037.

Timm,ia megapolitana Hedw.

Sandy soil, open places in prairie. Mitchell Co., McGregor 1999; Eiley
Co., M. A. Carleton (KSC).

308 The University Science Bulletin

Tortella humilis (Hedw.) Jennings.

On limestone rocks, moist soil in woods. Bourbon Co., McGregor 1777;
Cherokee Co., McGregor 1629, 1652; Linn Co., McGregor 1954.

Tortilla pagorum (Milde) De Not.

Bark of elm trees. Cherokee Co., McGregor 1021 ; Douglas Co., Mc-
Gregor 2655; Wilson Co., L. J. Gier, reported by Gier (1949), specimen in
herbarium at William Jewell College, Liberty, Missouri.

Weisia viridula Hedw.

This is apparently one of the most common mosses in the state. I have
found it in nearly every habitat I have visited. Barber Co., W. H. Horr 3115;
Butler Co., McGregor 2245; Chase Co., McGregor 1811, 1815; Chautauqua
Co.. McGregor 1195; Cherokee Co., McGregor 1026, 1032, 1648; Clark Co.,
McGregor 2208; Clay Co., McGregor 2073; Cloud Co., McGregor 2067; Cow-
ley Co., McGregor 2243, W. H. Horr 3167; Crawford Co., Holland lOO (KSC),
Gier 730 (KSTCP) ; Douglas Co., McGregor 1164, Thompson March 14, 1948;
Ford Co., McGregor 2211; Geary Co., F. C. Gates 18666 (KSC); Jewell Co.,
McGregor 2089; Johnson Co., McGregor 1334; Kingman Co., McGregor 2219,
2220; Kiowa Co., McGregor 2212; Leavenworth Co., McGregor 2260, 2262;
Lincoln Co., McGregor 1988; Meade Co., McGregor 2203; Montgomery Co.,
McGregor 1175, 2396; Nemaha Co., McGregor 2020; Ottawa Co., F. C. Gates
19086 (KSC), McGregor 1317, 1319; Phillips Co., McGregor 2097; Pratt
Co., McGregor 2218; Rawlins Co., McGregor 2135; Reno Co., McGregor
2222; Republic Co., McGregor 2086; Saline Co., McGregor 1488; Scott Co.,
McGregor 2163, 2174; Shawnee Co., McGregor 1230; Smith Co., McGregor
2093; Sumner Co., McGregor 2239; Washington Co., McGregor 2047; Wilson
Co., McGregor 1277.

EXCLUDED LIST

The following is a list of mosses which have been reported for the state
but which I am listing as excluded species. Unless otherwise indicated each
is excluded because of a lack of specimen evidence. Others are excluded for
reason given with the entry. If anyone knows of specimen evidence for any
of the following species I would appreciate receiving such information. Many
of them undoubtedly are present in Kansas.

Amblystegiella subtilis (Hedw.) Loeske.

Amblystegium carioaum (Sull.) This name does not occur in Grout's Moss

Flora of North America. The authority for the name is also in doubt. I

have found no one who has ever heard of this name.
Amblystegium compactum (C. M.) Aust.
Amblystegium radicale Bry. Eur. It is doubtful as to what this might have

been. It was mentioned in an early paper by Smyth but he omitted it in

hia paper of 1911. Probably it was A. varium.

Archidium hallii Aust.

Astomum crispum Hampe. Grout (1928-1940) says this plant does not occur
in North America. Entry might have been Astomum muhlenbergianum
(Sw.) Grout.

McGregor: Studies on Kansas Mosses I 309

Barbula convoluta Hedw.

Brachythecium acutum (Mitt.) Sull.

Brachythecium flagellare (Hedw.) Jennings.

Brachythecium rutahuhim (L.) Bry. Eur.

Brothera leana (Sull.) C. M.

Campylium polygamum (Bry. Eur.) Bryhn.

Campylium stellatum (Schreb. Hedw.) Lang, and C. Jens.

Climacium dendoides (L.) Web. and Mohr.

Dicranella rufescens (Dicks. Sm.) Schimp.

Dicranum fucescens Turner.

Dicranum undulatuni Turn. Is synonym of Dicranium bonjeani De Not. I

have not found specimen evidence.
Didymodon recurvirostris (Hedw.) Jenn.
Ditrichum lineare (Sw.) Lindb.
Ditrichum pusillum (Hedw.) E. G. Britton.
Drepanocladus aduncus (Hedw.) Warnst.
Entodon sullivantii (C. M.) Lindb.

Ephemerum crassinervhmi (Schwaegr.) C. Mull. var. papillosum (Aust.) Ren.
and Card.

Eurhynchium, praelongum (Dill L.) Bryhn.

Eurhynchium, riiscijorm,e (Neck.) Milde.

Eurhynchium, strigosum (Hoffn.) Bry. Eur.

Fabronia ciliaris (Brid.) Brid.

Fissidens bryoides Hedw.

Fissidens bryoides Hedw. var. incurvus (Starke) Monkem.

Fissidens exiguus Sull.

Fissidens osmundoides Hedw.

Fontinalis dalicarlica Bry. Eur.

Grimmia alpicola Hedw. var. rivularis (Brid.) Broth.

Grimmia apocarpa (L.) Hedw. var. conferta (Funck.) Spreng.

Grimmia apocarpa (L.) Hedw. var. conjerta (Funck.) Spreng f.

obtusifolia (Bry. Eur.) Moenk.

Grimmia calyptrata Hook.

Grimmia olneyi Sull.

Grimmia pilijera Beauv.

Grimmia wrightii Aust.

Gymnostomum aeruginosum Sm.

Hygroamblystegium, fluviatile (Sw.) Loeske.

Hygroamblystegium noterophilum (Sull.) Wamst.

Hygrohypnum alspestre (Hedw.) Loeske.

Hypnum porphyrrhizum Lindb. and Hypnum prophyrrizon Lindb. Reported
by Reed (1894) and Smyth (1941) are not in Grout (1928-1940). I am in

310 The University Science Bulletin

doubt as to what they might have had though it was probably Amblyste-

gium varium (Hedw.) Lindl.
Hypnum vagans Drum.
Leptodictyum vacillans (Sull.) Broth.
Leskea polycarpa (Ehrh.) Hedw.
Mnium affine elatwm Bry. Eur. Andrews in Moss Flora of North America

discarded all varietal names of Mnium affine.
Mnium elatum Bry. Eur. Probably same as above.
Myrinia pulvinata (Wahlenb.) Schimp.
Orthotrichum anamalum Hedw.
Orthotrichum cupulatum (Hoffm.) Brid.
Orthotrichum cupulatum (Hoffm.) Brid. var minus Sulliv.
Orthotrichum, speciosum, Nees.
Phascumi cuspidatum Hedw.
Philonotis longiseta (Rich) E. C. Britton.
Philonotis muhlenbergii (Schwaegr.) Brid.
Physcomitrium, acuminatum (Schleich.) Bry. Eur.
Plagiothecium sylvaticum (Huds. Brid.) Bry. Eur.
Pleuridium bolanderi C. Muell.
Pohlia annotina (Hedw.) Loeske.
Polytrichum pilifercum Hedw.
Pylaisia intricata Bry. Eur.
Bhacomitrium aciculare Brid.
Sphagnum, tabulare Sull.
Seligera pusilla (Hedw.) Bry. Eur.
Thuidium abietinum (L. Brid.) Bry. Eur.
Tortella caespitosa (Schwaegr.) Limpr.
Tortula m,ucronijolia Schwaegr.
Tricholepsis migrescens (Sw.) Grout
Trichostomum crispum Bruch.

BIBLIOGRAPHY OF KANSAS MOSSES

Andrews, A. LeRoy. Bryological notes I. Aschisma kansanum, a new species
with remarks upon the Genus. Torreya 15: 63-67. 1915.

, List of the North American Species of Sphagnum. Bryologist 43 No.

5, 132. 1940.

Britton, Elizabeth G. Contributions to American Biyology VII. A Re-
vision of the Genus Physcomitrium with Descriptions of Five New Species.
Bull. Torrey Bot. Club 21 : 189-208. 1894.

Cragin, F. W. Miscellaneous Notes. Bull. Washburn Lab. of Natural Science.
1: 83. 1885.

GiER, L. J. Flora of Crawford County (Kansas) State Park: Mosses. Trans.
Kan. Acad. Sci. 43: 119-120. 1940.

, Bryophytes of Kansas. Trans. Kan. Acad. Sci. 62 : No. 2, 58-65. 1949.

McGregor: Studies on Kansas Mosses I 311

Grout, A. J. Moss Flora of North America North of Mexico. 3 vols. Publ.
by the author, Newfane, Vt. 1928-1940.

. List of Mo.'^ses of North America North of Mexico. Bryologist. 43'

No. 5, 117-131. 1940.

LESQUERErx, Leo, and Thomas P. James. Manual of the Mosses of North
America. S. E. Cassino and Co. Boston. 1884.

Rau, Eugene A. First Contribution to the Knowledge of Kansas Mosses.
Bull. Washburn Lab. of Natural Science. 1 : 18. 1884.

-, Second Contribution to the Knowledge of Kansas Mosses. Bull.

Washburn Lab. of Natural Science: 1: 60-62. 1885a.

Third Contribution to the Knowledge of Kansas Mosses. Bull. Wash-

burn Lab. of Natural Science. 1 : 114. 1885b.

Fourth Contribution to the Knowledge of Kan.sas Mosses. Bull.

Washburn Lab. of Natural Science. 1 : 171-173. 1886.

Reed. Minnie. Kansas Mosses. Trans. Kan. Acad. Sci. U: 152-198. 1893-
1894.

Renauld, F., and J. Cardot. New Mosses of North America I. Botanical
Gazette 13: No. 8, 1888.

and , New Mosses of North America II. Botanical Gazette

U: No. 4, 91-99. 1889.

and , New Mosses of North America III. Botanical Gazette

15: 40-41. 1890.

and , Enumeration of the Kansas Mosses. Botanical Gazette

17: 81-85. 1892.

Smyth, B. B. Additions to the Flora of Kansas. Trans. Kan. Acad. Science.
12: 115-119. 1890.

, Additions the Flora of Kansas. Trans. Kan. Acad. Science 13: 102.

1891-1892.

-, Additions to the Plants of Kansas. Trans. Kan. Acad. Science 14'-

133-134. 1893-1894.

and L. C. Smyth. Catalogue of the Flora of Kansas Part I. Ttans.

Kan. Acad. Sci. ^3 and ^4: 280-287. 1911.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. II] March 20, 1950 [No. 8

Type Localities of Mexican Reptiles and Amphibians

BY

HoBART M. Smith and Edward H. Taylor,

University of Illinois, Department of Zoology, and
University of Kansas, Department of Zoology.

Abstract: This paper contains a list of the type localities of all the speci< :
of reptiles and amphibians at present recognized as belonging in the Mexica]
fauna. Type localities are also given for all species recognized as synonyms
of those forms.

The following list includes the type localities of all forms of rep-
tiles and amphibians at present recognized as occurring in Mexico
whether the type localities are in Mexico or elsewhere. The locali-
ties are arranged in alphabetical order throughout; the genera and
species are likewise arranged alphabetically within the eight groups
(in the order listed) of caecilians, salamanders, anurans, turtles,
amphisbaenians, lizards, snakes and crocodilians. The group to which
each name belongs is indicated by the prefixes g, u, a, t, am, s, o
and c, respectively. A number of forms mentioned in the current
and pending check lists, on which this list is based (see Smith and
Taylor, 1945, 1948, 1950) but not yet definitely known to occur in
Mexico, have been included. All synonyms noted in the lists men-
tioned are likewise included. Recognized forms are cited by the
name currently used printed in italicized bold-faced type; unrecog-
nized synonyms are cited by their original name, unmodified, and
printed in italics.

Compilation of the list has emphasized the surprising lack of
and the need for restriction of type localities, many of which are
currently expressed as "Mexico" or some other equally useless gen-
erality, or as a series of localities, none of which has been selected
as the type locality. We have made an attempt to correct the lack

(313)

314 The University Science Bulletin

of preciseness in statement of type locality as much as possible, by
restricting as many type localities as the present state of knowledge
will permit. We have restricted type localities only to localities
from which specimens are known to have been taken, and/or to
those at which there is a reasonable possibility the type or types
may have been secured.

We have omitted authors' names from scientific names in this list,
inasmuch as a precise reference to the form to which we refer (to-
gether with author and location of original description) may be
found under the name cited in the check lists mentioned. Inasmuch
as the validity of our restrictions of type locality of numerous names
hinges upon the clarity of bibliographic reference accompanying use
of those names, we wish to reiterate that: the bibliographic
reference we intend to accompany each name cited herein
whose type locality is designated "by present restriction" is
to be found under that same name (unless otherwise indi-
cated) in our check lists of 1945, 1948 and 1950 (see Smith and
Taylor, those dates). Some names * are not to be found in these
check lists, having been proposed after the check lists were in press.
They may be found in the articles cited in the bibliography.

EASTERN HEMISPHERE

CEYLON
No definite locality

t Caretta caretta gigas

EGYPT
Cairo

s Hemidactylus turcicus turcicus (by present restriction)

INDIA

Malabar

t Chelonia dussumieri
Vizagapatam

o Typhlops braminus

JAVA

No definite locality

s Hemidactylus frenatus

* Thorius miiiittinsimus. Thoriiis mocdougalli. Ptychohyla bogerti, Plectrohyla brachycephala,
Hyla proboscidpa, Hyla dalquesti, Rana moorei, R. pipiens auslricola, Arizona rlegans cbtirnata,
A. e. expolita, A. e. pacata, A. e. philipi, Crotalus basiliscus oaxacus, C. gloydi lautus, Ficimia
publia taylori, Hypsiglena ochrorln/ncha klauberi, H. o. texana, H. o. tortugaerisis, H. nnaondaris,
Micrunu nigrocinctus wagneri, Phyllorhynchus browni fortitus, Pituophis catenifer bimaris,
P. c. coronatis, P. r. fiiliginosus, P. c. int/ulanus, P, deppei brcvilineata, Pseudoficimia hiltoni,
Rhadinaea marcellae, Salvadora Iiexalepis klauberi, Schmidtophis rubriventris, Tantilla shawi,
Thamnophis melanogaster linearis, T. sumichrasti salvini, and Toluca amphisticha.

Mexican Reptiles and Amphibians 315

PHILIPPINE ISLANDS
Manila

t Lepidochelys olivacea
s Per opus mutilatus

PINE ISLAND

No definite locality
o Pelamis platurus

RED SEA

No definite locality

t Caretta bissa

SICILY
Palermo

t Dermochelys coriacea (by present restriction)

STRAITS SETTLEMENTS
Singapore

t Eretmochelys squamata (by restriction)

WESTERN HEMISPHERE

No definite locality

s Anolis damulus

s Anolis impetigosus

s Phyllodactylus mentalis

3 Phyllodactylus pidcher

o Dipsas bertholdi

o Erythrolamprus aesculapii

o Tropidodipsas annidifera

o Typhlops longissimus

ATLANTIC OCEAN

ASCENSION ISLAND

No definite locality

t Chelonia maculosa (by present restriction)
t Chelonia mydas

BERMUDA ISLANDS

No definite locality

t Caretta caretta caretta (by present restriction)

t Chelonia lachrymata (by present restriction)

t Chelonia pseudo-caretta (by present restriction)

t Chelonia pseudo-mydas (by present restriction)

t Chelonia virgala (by present restriction)

t Eretmochelys imbricata (by present restriction)

'616 The University Science Bulletin

CENTRAL AMERICA

BRITISH HONDURAS
Belize

t Dermatemys abnormis (by present restriction)

s Anolis ustus

s Coleonyx elegans elegans

o Elaphe triaspis (by present restriction)

o Herpetodryas incertus

o Leptodeira annulata polysticta (by restriction)

o Micrurus afRnis alienus (by present restriction)

0 Micrurus affinis stantoni

o Thamnophis sumichrasti praeocularis (by present restric-
tion)

o Tretanorliinus nigroluteus lateralis (by present restriction)

c Alligator lacordairei
Cohune Ridge (20 miles southeast of Benque Viejo)

s Sceloporus lundelli lundelli
Double Falls (west of Stann Creek)

o Pliocercus elapoides semicinctus
St. George Island (near Belize)

s Aristelliger georgeensis

COSTA RICA

No definite locality

s Mabuia alliacea
Arriba

a Bufo coccifer

s Anolis nannodes (by present restriction)
Bebedero

a Hyla underwoodi
Palmar

s Anolis capita (by present restriction)

s Leiolopisma cherriei cherriei
Palomo (Valle de Orosi)

a Anotheca coronata
San Jose

a Centrolenella Aeischmanni
San Mateo

s Cnemidophorus alfaronis

EL SALVADOR
Izalco, Volcan

s Leiolopisma assatum assatum

o Imantodes gemmistratus

La Union

t Geoemyda pulcherrima incisa

s Ctenosaura complcta (by present restriction)

s Sceloporus fulims

o Loxocemus bicolor

o Plastoseryx brunrn (by present restriction)

o Stenorrhina freminvillii lactea (by present restriction)

Mexican Reptiles and Amphibians 317

GUATEMALA

No definite locality

o Agkistrodon bilineatus (Pacific coast of Guatemala)

o Amastridium sapperi

o Pliocercus elapoides diastemus (plateau of)

o Tantilla jani

Alta Ver.\paz

No definite locality

t Claudius pictus (by present restriction)

o Petalognathus multijasciatus (by present restriction)
Coban

a Agalychnis moreletii

a Hyla holochlora

t Cinosternum cobanum (by present restriction)

s Anolis petersii biidttatiis (by present restriction)

s Anolis stimrti

s Sceloporus malachiticus taeniocnemis (by present restric-
tion)

o Bothriechis nummifera notata

o Bothrops mexicanus

o Bothrops nigroviridis aurifer

o Drymobius chloroticus

o Micrurus afhnis apiatus (by present restriction)

o Thaninophis suniichrasti fulvus (by present restriction)
Cubilguitz

o Leptognathus cuculliceps (by present restriction)

o Leptognathus semicinctus (by present restriction)

o Thamnophis sumichrasti salvini
Panzos, Rio Polochic

t Chelydra rossignonii

s Ameiva f estiva edwardsii (Santa Maria de, by present rcslric-
tion)

s Anolis copei

s Anolis limifrons rodriguezii

o Bothrops nasutus
Samac, Finca (6 kilometers west of Coban)

s Anolis petersii (by present restriction)
Volcan, Finca (49 kilometers east of Coban)

s Xenosaurus rackhami

Baja Verapaz

No definite locality

s Anolis carneus
San Geronimo

o Pliocercus aequalis

318 The University Science Bulletin

El Peten

Flores

o Coniophanes quinquevittatus (by present restriction)

o Tantilla moesta (by present restriction)
Ixpuc Aguada (north of La Libertad)

a Hyla loquax
La Libertad

t Geoemyda areolata (by present restriction)

t Gymnodactylus coleonyx (by present restriction)

s Gymnodactylus scapularis (by present restriction)

o Thalerophis occidentalis praestans (by present restriction) *
Peten, Laguna de

c Crocodylus moreletii
Piedras Negras

a Eleutherodactylus conspicuus

o Dendropliidion vinitor

o Scap/tiodontophis carpicinctus

o Tantillita lintoni

o Xenodon mexicanus
Rio de la Pasion

s Lepidophyma fLavomaculatum flavomaculatum (by present
restriction)
Santa Teresa, 2 miles north of

s Anolis humilis uniformis (by present restriction)

s Anolis ruthveni

ESCUINTLA

Escuintla

s Corythophanes percarinatus

o Thamnophis sumichrasti cerebrosus

Guatemala

Guatemala

o Geophis nasalis

HUEHUETBNANGO

Canibal, Finca (3,000 ft.)

s Sceloporus melanorhinus stuarti

Rbtaluleu

Huamuchil

t Staurotyjms biporcatus

t Staurotypus salvinii
Retaluleu

o Peropodum guatemalensis

o Ungaliophis continentalis

Sacatepbiquez

Antigua, Volcan

s Sceloporus squamosus (by present restriction)

• Formerly Leptophis occidentalis praestans.

Mexican Reptiles and Amphibians 319

SOLOLA

Atitlan

s Celestas atitlanensis

o Drymobius margaritiferus occidentalis (Volcan)

o Leptognathus sexscutatus

o Tropidodipsas Rscheri (Volcan) (by present restriction)

o Virginia fasciata (by present restriction)
San Agustin, 610 m. (near Volcan de Atitlan)

s Sceloporus malacluticus acanthinus

o Bothrops afjinis

o Bothrops bicolor

SUCHITEPEQUEZ

Finca El Cipres, Volcan Zunil

o Adelphicos quadrivirgatus sargii (by present restriction)

o Micrurus latifasciatus

o Micrurus nigrocinctus zunilensis

Mazatenango

s Akleistops guatemalensis
s Ameiva undulata parva (by restriction)
s Lepidophyma smithii smitliii (by restriction)
o Leptognathus bernoullii (Hacienda de Chitalon)
o Tropidodipsas sartorii annulatus (Hacienda de Chitalon, by
present restriction)

Rio Nagualate

t Chelonia agassizii
t Emys grayi
t Pseudemys umbra
c Crocodilus pacificus

TOTONICAPAM

Totonicapam

o Bothrops godmani

YZABAL

Yzabal

o Spilotes salvini (by present restriction)

Zac-Apa

Rio Motagua, Valley of (10 kilometers northeast of Zacapa)
s Cnemidophorus motaguae

HONDURAS

No definite locality

t Emys venusta

t Cristasaura mitrella

o Coniophanes fissidens punctigularis

o Microphis quinqueliniatus

o Pseustes poecilonotus poecilonotus
Lancetilla

s Eumeces schmidti

320 The University Science Bulletin

Portillo Grande (Yoro)

o Adelphicos qimdrivirgatus visoninus (by present restriction)
Tela

s Ctenosaura similis similis (by restriction)

NICARAGUA

No definite locality

a Hyla ebraccata

s Cnemidophorus decemlineatus
Greytown

s Basiliscus nuchalis

s Draconura bivittata (by present restriction)
Laguna Island

o Hypsiglena torquata
Rama

s Gonatodes fuscus (by present restriction)
Recero

a Leptodactylus melanonotus (by present restriction)

PANAMA
No definite locality

s Anolis panamensis
Chiriqui, Rio (in vicinity of Bocas del Toro)

a Bufo simus
Obispo

o Bothrops atrox asper
Veragua

s Splmerodactylus lineolatus

NORTH AMERICA

MEXICO
State unknown

u Magnadigita sulcata

u Spelcrpes pnnciatum

a Eleutherodactylus longipes

a Hyla crassa

a Hyla plicata

t Cinosicrnum brcvigulare (Playa Vicente)

s Anolis baccatus

s Anolis cumingii

s Anolis guniherii

s Anolis heliactin

s Anolis nietallicus

s Anolis schiedii

s Urosaurus irregularis (highlands)

o Anomalepis mexicana

o Dipsas brevis

o Dipsas dimidiatus

o Erythrolamprus guentheri

o Geophis chalybeus

Mexican Reptiles and Amphibians 321

o Geophis rostralis

o Geophis sieboldi (?)

o Helicops septemvittata

o Rhadinaea taeniata

o Sonora aeqiuilis

o Tantilla deppei

o Tantilla mexicana

o Typfdops psittacus

Baja California

No specific locality

s Phyllodactylus tuberculosus ( Calif ornien)
Mainland localities

Agua Calieute (Cape region)

o Crotalus lucasensis
Agua Verde Bay

s Callisaurv^ plasticus
Alcatraz

o Lampropeltis zonata agalma
Ballenas Bay

s Callisaurus draconoides crinitus
Comondu

s Eumeces lagunensis (on trail between Loreto and; neo-
type locality)

s Petrosaurus repens

o Sonora mosaueri
Ensenada

s Cnemidophorus stejnegeri

o Lichanura myriolepis (by present restriction)

o Lichanura roseofusca roseofusca (by present restriction)
Estero Salina

o Chilomeniscus stramineus esterensis
Gardners Laguna, Salton River

s Callisaurus draconoides gabhii (by present restriction)
La Paz

s Coleonyx variegatus peninsularis

s Crotaphytus copeii

s Ufa stansburiana elegans

o Chilomeniscus stramineus jasciatus

o Leptotyphlops humilis slevini
Las Palmas

s Crotaphytus fasciolatus
Los Dolores

t Pseudemys scripta nebulosa
Puerto Escondido

o Hypsiglena slevini
San Bartolome Bay

s Cnemidophorus bartolomas

s Phrynosoma coronatum jamesi

s Uta parva

20—90

322 The University Science Bulletin

San Fernando Mision (between San Ignacio and)

o Phyllorhynchus decurtatus decurtatus (by present re-
striction)
San Francisquito (Sierra de la Laguna)

s Eumeces lagunensis

s Xantusia gilberti
San Francisquito Bay

s Sauromalus australis
San Ignacio

o Thatnnophis digueti (by restriction)
San Jorge, 2 miles east of

o Sonora bancroftae
San Jose del Cabo

s Dipsosaurus dorsalis lucasensis

o Lampropeltis nitida {^= L. getulus nitida)
San Lucas, Cape

a Hyia curta

a Hyla regilla laticeps

a Scaphiopus varians
am Bipes biporus

s Callisaurus drnconoides draconoides (by present restric-
tion)

s Cnemidophorus hyperythrus hyperythrus

s Cnemidophorus maximus

s Ctenosaura hemilopha

s Ctenosaura interrupta (by present- restriction)

s Petrosaurus thalassinus

s Phrynosoma coronatum coronatum (by present restric-
tion)

s Phyllodactylus unctus

s Phyllodactylus xanti

s Sceloporus magister zosteromus

s Urosaurus nigricaudus

o Chilomeniscus stramineus stramineus

o Cro talus enyo

0 Crotalus mitchellii mitchelUi

o Hypsiglena ochrorhynclia ochrorhyncha

o Lampropeltis getulus conjuncta

o Lichanura trivirgata

o Masticoplus aurigulus

o Natrix valida celaeno

o Pituophis catenifer vertebralis (by present restriction)

o Pityophis haeniatois

o Salvadora hexalepis klauberi

0 Tantilla planiceps (by present restriction)

o Trimorphodon lyroplmnes

o Tropidonotus tephropleura
San Pedro Martir Mountains

s Urosaurus microscutatus

o Crotalvs goldmani (El Pinon, 5,300 ft.)

Mexican Reptiles and Amphibians 323

o Thamnophis ordinoides hueyi (=r T. elegans hueyi)
(Arroyo Encantado)
San Quintin

s Phrynosoma nelsoni
San Quintin Bay

s Sceloporus magister ru/idorsum
Santa Gertrudis, near El Arco

o Pituophis catenifer bimaris
Santa Rosalia

s Sceloporus digueti

o Elaphe rosaliae

o Hypsiglena ochrorhyncha venusta (by present restriction)
Santo Domingo

s Callisaurus rhodostictus (by present restriction)

o Arizona elegans pacata
Sierra San Lazaro

s Sceloporus orcutti licki
Tecate

s Phrynosoma ochoterenai
Todos Santos

s Elgaria paucicarinata
Insular localities

Angel de la Guarda

s Callisaurus draconoides splendidus

s Crotaphytus insularis

s Sauromalus hispidus

o Crotalus atrox elegans
Carmen

s Callisaurus draconoides carmenensis

s Cnemidophorus hyperythrus caeruleus

s Cnemidophorus vandenburghi

s Dipsosaurus carmenensis

s Uta mannophorus
Cedros

s Cnemidophorus hyperythrus beldingi

s Cnemidophorus tigris multiscutatus

s Cnemidophorus labialis

s Elgaria cedrosensis

s Phrynosoma cerroense

s Phrynosoma schmidti

s Uta concinna

o Crotalus exsul

o Pituopfiis catenifer insulanus
Ceralvo

s Cnemidophorus ceralbensis

s Ctenosaura insulana

s Sat or grandaevus
Los Coronados

u Batrachoseps attenuatus leucopus (North Island)

s Elgaria multicarinata nana (South Island)

324 The University Science Bulletin

o Hypsiglena ochrorhyncha klauberi (South Island)

o Pituophis catenifer coronalis (South Island)
Espiritu Santo

s Sauromalus ater (by present restriction)

s Verticaria espiritensis
Isla Parti da

s Cnemidophorus dickersonae (near Angel de la Guarda)

o Chilomeniscus punctatissimus (near Espiritu Santo)

o Masticophis barbouri (near Espiritu Santo)
Mejia

s Streptosaurus slevini
Monserrate

s Cnemidophorus hyperythrus pictus

s Sauromalus slevini (southern end)

s Sceloporus magister monserratensis
Sal Si Puedes

s Cnemidophorus tigris canus
San Benito

s Uta stellata
San Francisco

s Verticaria jranciscensis
San Geronimo

s Anniella geronimensis
San Jose

s Cnemidophorus hyperythrus danheimae

s Cnemidophorus tigris celeripes
San Marcos

s Cnemidophorus hyperythrus schmidti
San Martin

s Gerrhonotus scincicauda ignavus

s Uta martinensis

o Pituophis catenifer fuliginatus
Santa Catalina

s Cnemidophorus catalinensis

s Dipsosaurus catalinensis

s Sauromalus klauberi

s Sceloporus magister lineatulus

s Uta squamata

o Lampropeltis catalinensis
Santa Cruz

s Sator angustus
Santa Inez, South

s Coleonyx variegatus slevini
Santa Margarita

s Cnemidophorus tigris rubidus

o Zamenis lateralis fuliginosus
Todos Santos, South

o Diadophis amabilis anthonyi

o Lampropeltis zonata herrerae

Mexican Reptiles and Amphibians 325

Tortuga

o Crotalus tortugensis

o Hypsiglena ochrorhyncha tortugaensis

Campechb

No definite locality

s Eumeces schivartzei (a small island in the Laguna de Ter-
minos)
Balchacaj

s Ctenosaura (Cachryx) annectens (by present restriction)

s Enyaliosaurus erythramelas (by present restriction)

o Imantodes splendidus luciodorsus
Encarnacion

a Hypopachus cuneus nigroreticulatus
Panlao

s Anolis sagrei mayensis
Tuxpena

o Leptodeira yucatanensis malleisi

Chiapas

Chicharros

u Bolitoglossa flaviventris

o Geophis cancellatus
Ciudad de las Casas, 11 kilometers southeast of, 2,300 m.

s Barisia moreleti temporalis
Conjab, between San Bartolome and Comitan, 5,300 ft.

s Sceloporus prezygus
Finca Germania, 4 miles northeast of Huixtla, 3,300 ft.

o Micrunis nigrocinctus wagneri
La Esperanza, near Escuintla

u Bolitoglossa occidentalis

a Acrodytes niodesta (Cruz de Piedra)

a Bufo angustipes

a Microbatrachylus imitator

a Syrrhophus rubrimaculatus

s Sceloporus guentheri (by present restriction)

o Drymarchon corais unicolor

o Scaphiodontophis albonuchalis
La Libertad

s Ameiva undulata thomasi
Male, Cerro, 3,200 m.

s Gerrhonotus liocephalus austrinus
Mineral de Santa Fe

s Eumeces rovirosae
Ovando, Cerro

u Chiropterotriton xolocalcae

u Magnadigita nigroflavescens

a Eleutherodactylus matudai

a Hyla rozellae (Salto de Agua)

a Microbatrachylus montanus

326 The University Science Bulletin

a Plectrohyla matudai

a Plectrohyla sagorum

o Elaphe flavirufa niatudai (Salto de Agua)

o Micrurus nigrocinctus ovandoensis (Salto de Agua)

o Rhadinaea lachrymans (by present restriction)
Palenque

u Bolitoglossa moreleti

s Ameiva undulata stuarti

s Celestas rozellae

o Coniophanes bipunctatus biseriatus

o Thamnophis rozellae
Piedra Parada

s Anolis biporcatus (by present restriction)
Piedras Negras, Guatemala, across Rio Usumacinta from

s Ameiva undulata hartwegi
San Cristobal

o Adelphicos veraepacis nigrilatus
San Ricardo

a Hypopachus maculatus
Santa Rosa, near Comitan

s Abronia ochoterenai
Siltepec, 16 miles south of, 2,300 m.

s Barisia moreleti rafaeli
Tacana, Volcan de

s Abronia matudai
Tapachula

a Hyla beltrani

a Hyla robertmertensi
Tonala

a Syrrhophus nebulosus

o Conophis pulcher plagosus

o Conophis pulcher similis (by present restriction)

o Tantillita brevissima

c Alligator Chiapasius
Tuxtla Gutierrez

t Kinosternon abaxillare

s Sceloporus carinatus

Chihuahua

No definite locality

s Barisia levicollis * (by present restriction)

o Trimorphodon vilkinsonii
Batopilas

s Urosaurus unicus (?)

o Procinura aemula
Batosegachic

o Micrurus diastema distans

Originally stated "Mexican Boundary," the type locality cannot reasonably be restricted
further than "Chihuahua" at the present time, inasmuch as no specimens are yet known from
localities near the U. S. boundary in that state.

Mexican Reptiles and Amphibians 327

Casas Grandes

o Arizona elegans expolita

Chihuahua

a Bufo insidior

s Cnemidophorus gularis gularis obsoletus

s Cnemidophorus gularis gularis verus

a Cnemidophorus sackii scalaris

s Cnemidophorus sexlineatus tigris Cope (nee Baird and
Girard)

s Urosaurus ornatus caeruleus (30 miles north of)
Colonia Garcia

o Thamnophis ordinoides errans (= Thamnophis errans)
Guasaremos, Rio Mayo

o Tantilla yaquia
Juarez, 15 miles south of

s Holbrookia hunkeri
Mojarachic

u Ambystoma Huvinatum

u Ambystoma rosaceum

a Eleutherodactylus tarahumaraensis

s Elgaria kingii kingii (by present restriction)

s Gerrhonotus multijasciatus (by present restriction)

o Crotalus semicornutus

o Lampropeltis knoblochi
Presidio del Norte

o Churchillia bellona
San Buenaventura, 11 miles south of

o Masticophis flagellum lineatulus
Sierra Madre, lat. 26°6' N., long. 106°50' W.

u Axolotes maculaia
Yoquivo

a Rana tarahumarae (by present restriction)

COAHUILA

Agua Nueva

s Cnemidophorus semifasciatus
Arteaga

s Sceloporus jarrovii oberon
Buena Vista

o Pituophis deppei jani
Castanuelas

s Holbrookia maculata dickersonae
Cuatro Cienegas

t Terrapene coahuila
El Salado, San Luis Potosi, 30 miles north of

s Sceloporus cautus
Monclova

o Crotalus (tigris) palmeri

o Rhinocheilus lecontei tessellatus

328 The University Science Bulletin

Nazas, Rio

a Scaphiopus rectifrenis
PaiTas

s Cnemidophorus variolosus

o Thamnophis angustirostris
Patos

s Sceloporus ornatus ornatus
Rinconada

o Thamnophis eques cyrtopsis
San Pedro (de las Colonias)

s Sceloporus ornatus caeruleus (5 miles south of)

s Unta exsul (sand dunes 12 miles north of)
Sierra Guadalupe

s Barisia imbricata ciliaris

COLIMA

No specific locality

s Phrynosoma orbiculare dugesii

s Sceloporus dugesii dugesii

o Geophis semiannulatus

o Leptotyphlops dugesii
Colima

a Agalychnis dacnicolor

a Bujo argillaceous (by present restriction)

s Cnemidophorus communis copei (by present restriction)

s Cnemidophorus deppii lineatissimus (by present restriction)

s Cnemidophorus sackii communis (by present restriction)

s C tenosaura brevirostris

s Ctenosaura pectinata (by restriction)

s Eumeces colimensis

s Phrynosoma asio (by present restriction)

s Sceloporus horridus oligoporus

s Sceloporus pyrocephalus

s Sceloporus utiformis

s Urosaurus bicarinatus tuberculatus

o Crotalus basiliscus basiliscus (by present restriction)

o Elaps epistema (by present restriction)

o Elaps fulvuis hypostema (by present restriction)

o Micrurus diastema diastema (by present restriction)

o Pseudoficimia frontalis (by present restriction)

o Pseudoficiynia pulchra (by present restriction)
Comala

o Tropidodipsas occidentalis
Manzanillo

s Ameiva undulata sinistra

s Anolis schmidti
Paso del Rio, Hacienda

a Syrrhophus modestus

a Coleonyx elegans nemoralis

o Dipsas gaigeae

o Leptotyphlops phenops bakewelli

Mexican Reptiles and Amphibians 329

Santiago

s Leiolopisma assatum taylori

Revillagigedos Archipelago

s Urosaurus auriculatus (Socorro Island)
s Urosaurus clarionensis (Clarion Island)
o Hypsiglena unaocularis (Clarion Island)
o Masticophis anthonyi (Clarion Island)

DisTRiTO Federal

Coyoacan

a Hyla eximia (bj^ present restriction)

a Scaphiopus dugesii (by present restriction)

a Scaphiopus multiplicatus (by present restriction)

o Crotalus triseriatus anahuacus (by present restriction)

o Oxyrhina de filippii (by present restriction)

o Toluca lineata lineata (by present restriction)

Desierto de los Leones
a Hyla lafrentzi

Laguna Santa Isabel, near Guadalupe Hidalgo
u Ambystoma velasci
u Siredon tigrina

Magdalena

s Scincus ventralis (by present restriction)

Mexico (City)

a Rana adtrita (by present restriction)
a Rana montezumae

s Agania torquata Peale and Greene (by present restriction)
s Barisia itnbricata imbricata (by present restriction)
s Gerrhonolus lichenigerus (by present restriction)
s Phrynosoma orbiculare orbiculare (by present restriction)
s Phrynosoma wiegmanni (by present restriction)
s Sceloporus grammicus microlepidotus (by present restric-
tion)
s Sceloporus scalaris scalaris (by present restriction)
s Sceloporus torquatus torquatus (by present restriction)
s Tapaya orbicularis longicaudatTis (by present restriction)
o Diadophis dugesii (Potreros de Balbuena, near)
o Eutaenia flavilabris (by present restriction)
o Eutaenia insigniarurn (Chapultepec)
o Eutaenia macrostemma
o Geophis bicolor (?)
o Rhadinaea laureata
o Thamnophis scalaris scaliger (by present restriction)

San Diego

s Gerrhonotus olivacev^ (by present restriction)

Serrania de las Cruces, Manantial de los Axolotes
u Rhyacosiredon altamirani

Xochimilco

u Siredon mexicanum (by present restriction)
a Bufo anomalus (by present restriction)

330 The University Science Bulletin

a Bufo compactilis compactilis (by present restriction)
a Bufo levifrons (by present restriction)
a Bujo mexicanus (by present restriction)

o Thamnophis melanogaster melanogaster (by present restric-
tion)
o Tropidonohis mesomelanus (by present restriction)

DURANGO

No specific locality

s Phrynosoma douglassii brachycercum

o Natrix valida valida
Pedricena

s Sceloporus lineolateralis (6 miles northeast of)

s Sceloporus maculosus (14 miles northeast of)
Ventanas

a Rana pustulosa

s Coleonyx fasciatus

Guanajuato

Acambaro

o Lampropcltis triangulum nelsoni (L. doliata nelsoni)

o Salvadora bairdii (by present restriction)
Guanajuato

a Bufo monksiae (by present restriction)

a Bufo occidentalis (by present restriction)

a Eleutherodactylus augusti

a Rana montezumae concolor (by present restriction)

a Syrrhophus guttilatus

t Cinosternum rostellum

s Eumeces callicephalus (by present restriction)

s Eumeces dugesii (by present restriction)

s Sceloporus jerrariperezi (by present restriction)

o Conopsis nasus (by present restriction)

o Epirhina tessellata (by present restriction)

o Eutaenia p^dchrilatus

o Hemigenius variabilis

o Hypsiglena ochrorhyncha janii (by present restriction)

o Masticophis taeniatus australis (by present restriction)

o Micrurus Mzingeri Rtzingeri (by present restriction)

o Oxyrhina (Exorhina) maculata (by present restriction)

o Tantilla bocourti (by present restriction)

o Thamnophis cyrtopsis cy elides (by present restriction)
Salamanca

o Thamnophis stejnegeri
Sierra de Santa Rosa

o Oreophis boulengeri
Tupataro

s Sceloporus torquatus melanogaster (by present restriction)

o Crotalus lugubris multimaculata (by present restriction)

o Crotalus polystictus (by present restriction)

Mexican Reptiles and Amphibians 331

Guerrero

Acapuico

t Kinosternon integrum (by present restriction)

s Anolis taylori (1 mile north of)

s Phyllodactylus magnatuberculatuti

a Sceloporus melanorhinus calligaster

o Conophis vittatus vittatus (Laguna Coyuca, by present re-
striction)

o Dryadophis melanolomus stuarti

o Imantodes gracillimus (by present restriction)

o Micrurus nuchalis taylori

o Salvadora lemniscata (by present restriction)

o Trimorphodon biscutatus semirutus
Agua del Obispo (between Rincon and Cajones)

a Centrolenella viridissima

a Hyla erythromma

a Hyla pinorum

a Hypopachus caprimimus

a Microbatrachylus minimus

a Ptychohyla adipoventris

a Rana sierramadrensis

a Tomodactylus albolabris

s Anolis dunni

s Anolis megapholidotus
Amula

a Tomodactylus amulae
Balsas, Rio

o Leptodeira guilleni
Buena Vista

o Tropidodipsas guerreroensis
Chilpancingo

a Hyla melanomma (7 miles east of)

s Sceloporus grammicus grammicus (by present restriction)

s Sceloporus pleurostictus Weigmann (by present restriction)

o Enulius unicolor (by present restriction)

o Leptotyphlops maximus

o Micrurus broumi (^ M. nigrocinctus browni)

o Salvadora intermedia intermedia

o Tantilla coronadoi
El Treinte

o Rhadinaea hesperia baileyi

o Tantilla martindelcampoi
Huajintlan (at Morelos border)

s Heloderma Hernandesii (by present restriction)

s Heloderma horridum (by present restriction)

o PseudoRcimia pulcherrima (by present restriction)

o Pseudoleptodeira latifasciata (by present restriction)
La Venta

a Acrodytes infiata

a Bufo gemmifer (El Limoncito, near)

o Coniophanes fissidens dispersus (El Limoncito, near)

332 The University Science Bulletin

Malinaltepec

a Hyla leonard-schultzei
Mazatlan

a Syrrhophus pipilans (9 miles south of)

s Eumeces ochoterenai

s Sceloporus ochoterenae (2 miles north of)
Mexcala

a Bufo perplexus
am Bipes canaliculatus (by present restriction)
am Chalcides sulcata (by present restriction)
am Chamaesaura propus (by present restriction)
am Lacerta lumbricoides (by present restriction)
am Lacerta mexicana (by present restriction)
Mezquititlan (north of Chilpancingo)

s Sceloporus gadovii
Omilteme

a Eleutherodactylus calcitrans (by restriction)

a Eleutherodactylus saltator

a Hyla arboricola (6 miles east of)

a Rana omilte^nana

s Abronia deppii (by present restriction)

s Anolis liogaster (7,600 ft.)

s Barisia gadovii gadovii

s Sceloporus formosus scitulus

s Sceloporus mucronatus omiltemanus (8,000 ft.)

s Sceloporus ruhriventris

o Agkistrodon hrowni

o Bothrops barbouri

o Crotalus omiltemanus

o Geophis omiltemana

o Rhadinaea aemula (by present restriction)

o Rhadinaea hesperia hesperia (by present restriction)

o Rhadinaea omiltemana

o Thamnophis scalaris godmani
Rincon

s Ameiva undulata dextra
Taxco

s Phyllodactylus bordai (6 miles north of)

o Sonera erythrura (10 miles south of)
Tecpan de Galeana
am Bipes tridactylus

o Geatractus tecpanecus
Tierra Colorada

s Anolis gadovii

s Phyllodactylus delcampi

s Phyllodactylus lanei

s Phyllodactylus magnus

s Sceloporus stejnegeri
Tixtla

o Ficimia ruspator (3 miles east of)

Mexican Reptiles and Amphibians 333

Hidalgo

Durango

u Cfiiropterotriton chondrostega

u Chiropterotriton mosaueri (7,200 ft.)

s Gaigeia gaigeae

o Ninia diademata plorator

o Rliadinaea crassa
El Chico

a Hyla rohertsorum (Parque Nacional)

s Abronia taeniata taeniata

s Eumeces lynxe lynxe

s Plestiodon bellii
El Pinalito

s Sceloporus jarrovii immucronatus (10 miles north of)
Guerrero (near Mineral del Monte)

u Chiropterotriton dimidiata

u Pseudoeurycea cephalica nianni
LaPlacita (south of Jacala), 7,000 ft.

a Tomodactylus macrotympanum

s Leiolopismd forbesorum
Santa Anita

u Pseudoeurycea cephalica rubrimembris (6 kilometers south
of)
Tianguistengo

u Chiropterotriton arborea

u Chiropterotriton terrestris (6 miles south of, 5,000 ft.)

a Eleutherodactylus hidalgoensis (4 miles west of)

a Hyla bronieliana
Zacualtipan

a Syrrhophus verrucipes (1,800 ft. lower than)

s Gaigeia sylvatica (7 miles north of)

o Geophis mutitorques

o Micrurus bernadi

o Storeria hidalgoensis

o Thamnophis phenax halophilus (7 miles north of)
Zimapan

s Phrynosoma boucardi (by present restriction)

Jalisco

Barranca Ibarra

s Ctenosaura parkeri
Chapala

a Rana megapoda

o Thamnophis melanogaster canescens
Cofradia

s Urosaurus gadovii
Corrientes, Cape

o Salvadora mexicana
Guadalajara

a Microhyla usta usta

a Rana trilobata {^ R. pipiens trilobata)

334 The University Science Bulletin

o Adelophis copei

o Cheilorhina viUarsii

o Clelia clelia imniaculata

o Coniophanes lateritius

o Conophis sumichrastii sumichrastii

o Crotalus jimenezii (by present restriction)

o Imantodes latistratus (by restriction)

o Sympholis Uppiens

o Tantilla calamarina

o Thamnophis subcarinata subcarinata (by restriction)

o Trimorphodon upsilon
La Cumbre de los Arrastrados /

s Sceloporus asper

s Sceloporus bulleri

s Sceloporus heterolepis (by present restriction)
Magdalena

s Sceloporus scalaris unicanthalis

0 Hypsiglena affinis (by present restriction)

o Rhadinaea hesperia hesperioides

o Sonora michoacanensis niutabilis
San Ramon

o Elaphe chlorosoma

Mexico

Asuncion

s Eumeces copei (10 miles southeast of)
Gavia, Hacienda de la

s Barisitt rudicollis (by present restriction)
Guadalupe, Rancho (14 kilometers east of San Martin)

u Ambystoma bombypellum

u Ambystoma schmidti
Ixtapan del Oro

a Eleutherodactylus bolivar i
Lake Lerma

u Siredon lermaensis

o Thamnophis melanogaster linearis
Rio Frio

u Rhyacosiredon leorae
San Juan Teotihuacan

o Elaphis pleurostictus (by present restriction)

o Pituophis deppei deppei (by present restriction)

o Pituophis depqtei pholidostictus (by present restriction)
San Martin

s Gerrhonotus adspersics (by present restriction)
Toluca

u Ambystoma granulosum (kilometer no. 74, about 12 miles
northwest of)

u Pseudoeurycea robertsi (Nevado de, 10,000 to 11,000 ft.)
Villa Victoria

u Rhyacosiredon rivularis (13 kilometers west of)

o Conopsis biserialis

Mexican Reptiles and Amphibians 335

Zempoala

s Sceloporus jarrovii sugillatus (Laguna no. 4)
Zumpango, Lake

u Ambystoma lacustris

MiCHOACAN

Apatzingan (de la Constitucion)

a Cystignathus microtis (by present restriction)

s Eumeces altamirani (by restriction)

o Micrurus diastema michoacanensis (by present restriction)

o Sonora michoacanensis michoacanensis (by present restric-
tion)
Cicio

o Geophis maculiferus
Cojumatlan

a Hyla smaragdina (6 kilometers east of)
El Mirador

u Ambystoma ordinarium (4 miles west of, near Puerto Hondo)
El Sabino (near Uruapan)

a Eleutherodactylus vocalis

o Leptodeira bressoni

o Leptodeira smithi

o Leptotyphlops bressoni

o Thamnopliis eques postremus
La Noria, near Zamora, Hacienda of D. Epifanio Jimenez

s Sceloporus dugesii intermedins

s Sceloporus westphalii
Morelia

u Ambystoma amblycephalum (15 kilometers west of)
Ovopeo, 1,000 ft.

s Enyaliosaurus clarki
Patzcuaro

u Bathysiredon dumerilii (Lake)

o Geophis petersii (by present restriction)

o Lampropeltis ruthveni
Quiroga

a Tomodactylus angustidigitorum
Tancitaro, 6,000 ft.

o Pituophis deppei brevilineata
Tangancicuaro

o Geophis dugesii
Temaxcal

o Thamnopliis vicinus
Tengohecho

o Erythrolamprus grammophrys
Uruapan

a Microbatrachylus hobartsmithi

o Trimorphodon fasciolata (Zararacua Falls)

336 The University Science Bulletin

MORELOS

Cuernavaca

a Hyla smithi

a Hypopachus alboventer (8 miles east of)

s Cnejnidophorus costatus (by present restriction)

s Cnemidophorus mexicanus balsas (by present restriction)

s Cnemidophorus sackii sackii (by restriction)

s Eumeces indubitus (kilometer 63)

s Sceloporus horridus horridus (by present restriction)

s Urosaurus bicarinatus bicarinatus (by present restriction)

Tepoztlan

a Hylella azteca

Tres Cumbres

s Seeloporus aeneus aeneus (by present restriction)

o Crotalus transversus

o Storeria storerioides (by present restriction)

Zempoala (Lakes of)

u Pseudoeurycea altamontana
u Rhyacosiredon zempoalaensis

Nayarit

Acaponeta

o Tantilla bogerti
Ixtlan

s CnemidopJiorus communis occidentalis (by present restriction)
Miramar

o Drymobius margaritiferus Rstulosus
Pedro Pablo, 2,500 ft.

t Terrapene nelsoni
San Bias

s Hemidactylus navarri

o Manolepis putnami

o Rhinocheilus lecontei antonii
Tepic

a Bufo nayaritensis

a Hyliola digueti

a Hypopachus ovis

a Leptodactylus occidentalis

s Eumeces parvulus

s Sceloporus horridus albiventris

s Sceloporus obscurus
Tres Marias Islands

s Cnemidophorus sackii mariarum

o Constrictor constrictor sigma (Maria Madre Island)

o Dryadophis melanolomus slevini (Maria Madre Island)

o Drymarchon corais cleofae (Maria Cleofa Island)

o Exelencophis nelsoni (Maria Madre Island)

o Lampropeltis triangulum schmidti (== L. doliata schmidti)

o Masticopliis flagellum lariolosus (Maria Magdalena Island)

o Thalerophis diplotropis forreri

Mexican Reptiles and Amphibians 337

NuBivo Leon

No exact locality

o Lampropeltis leonis

o Tantilla atriceps
Galeana

u Pseudoeurycea galennae (7,000 ft.)

a Syrrhophus smithi (15 miles west of)

o Crotalus triseriatus miquihuana (^ C. pricei miquihuana ;
Cerro Potosi)

o Rhadinaea montana (Ojo de Agua, near)

o Tantilla wilcoxi rubricata (15 miles southeast of)
Montemorelos

s Cnemidophorus gularis meeki
Monterrey

a Syrrhophus latodactylus (Huasteca Canon, 15 miles west of)

s Sceloporus cyanogenys
Pablillo, trail between Alamar and

s Sceloporus torquatus binocularis
Pesqueria Grande

s Cnemidophorus perplexus inornatus

s Cnemidophorus sackii octolineatus
Sabinas Hidalgo

s Sceloporus parvus parvus (hills 5 miles west of)

o Pituophis mexicanus
Santa Caterina

s Sceloporus couchii

o Nerodia couchii (by present restriction)

Oaxaca

No definite locality

a Eleutherodactylus rugulosus (Pacific region of the Isthmus

of Tehuantepec) *
s Abronia oaxacae

s Anolis boulengerianus (Isthmus of Tehuantepec)
s Anolis rubigenosiis
s Barisia imbricata planifrons
s Cnemidophorus unicolor (West Tehuantepec)
s Phrynosoma braconnieri
o Coluber oaxaca
o Enulius sumichrasti (Isthmus of Tehuantepec) *

o Lampropeltis triangidum oligozona (Tehuantepec = L. doliata

oligozona)
o Pseudoleptodeira discolor
o Salvadora bogerti (Tehuantepec)
Buena Vista

o Rhadinaea macdougalli

* More precise restriction is prohibited by the absence of definite records from southeastern
Oaxaca where the types undoubtedly were obtained. Very likely the types were collected near
Tapanatepec.

21—90

338 The University Science Bulletin

Cacoprieto

o Scaphiodontophis sumichrasti
Chihuitan

o Coniophanes piceivittis
o Symphimus leucostomus
Concordia, Cafetal, 600 m., between Puerto Angel and Salina Cruz
g Gymnopis multiplicata oaxacae
u Magnadigita macrinii
Cuicatlan

s Cnemidophorus sackii australis (by present restriction)

s Urosaurus bicarinatus nelsoni
Dondominguillo

s Ctenosaura multispinis
El Barrio

o Stenorrhina freminvillii apiata
Guichicovi

o Ficimia variegata (by present restriction)
Ixtepec (^ San Geronimo)

s Sceloporus edivardtaylori
Juchitan

t Geoemyda rubida

s Cnemidophorus lativittis

o Toluca lineata acuta
Lachiguiri, 7,100 ft.

6 Gaigeia dontomasi
La Gloria, north of Niltepec

a Eleutherodactylus macdougalli
Maquiltianguis

u Thorius macdougalli (Cerro de Humo)
Mazatlan

s Sphaerodactylus glaucus torquatus
Mixtequilla

o Tantilla depressa

o Tantilla striata
Niltapec

s Ficimia ramirez (1 league north)
Oaxaca

s Barisia gadovii levigata (valley of)

s Barisia viridiilava (by restriction)

s Cnemidophorus sackii bocourti (by present restriction)

s Eumeces brevirostris (by present restriction)

s Gerrhonotus bocourti (by present restriction)

s Gerrhonotus obscurus (by present restriction)

o Crotalus basiliscus oaxacus
Pluma Hidalgo

o Geophis sallaei (by present restriction)
Putla

s Anolis nebuloides
Quezaltepec

s Anolis milleri

Mexican Reptiles and Amphibians 339

Quiotepec

o Trimorphodon tau

Rio Astuta

a Plectrohyla brachycephala (at the foot of the Sierra Madre
between the Sierra Madre and the Sierra Atrevesado)
Rio Grande

a Ptychohyla bogerti
San Felipe, Cerro (15 kilometers northeast of Oaxaca)

u Bolitoglossa mexicana (by present restriction)

u Pseudoeurycea cochranae

11 Pseudoeurycea unguidentis

u Salamandra togata (by present restriction)

u Thorius narisovalis

u Thorius pulmonaris

a Eleutherodactylus mexicanus (by present restriction)

a Hyla hazelae

a Hyla robustofemora

a Microbatrachylus lineatissimus (7,000 ft.)

a Microbatrachylus oaxacae

o Crotalus gloydi (10,000 ft.)

o Thamnophis eburatus

o Toluca megalodon (summit of)
San Jose Manteca (5 kilometers from San Carlos Yautepec)

s Gaigeia radula
San Juan del Rio

t Rhinoclemmys mexicana
San Juan Guivini

o Toluca conica
San Luis, Cerro (15 kilometers northeast of Oaxaca)

u Pseudoeurycea smithi

s Sceloporus spinosus caeruleopunctatus
San Mateo del Mar

a Microhyla usta gadovii

t Cinosternum triliratum (by present restriction)

t Kinosternon cruentatum cruentatum (by present restriction)

t Kinostemum mexicanum (by present restriction)
Santa Efigenia

a Hylella sumichrasti

a Syrrhophus leprus

i Claudius severus

t Staurotypus marmoratus (by present restriction)
Tapanatepec

a Bufo canaliferus (by present restriction)

a Cystignathus perlaevis

a Hylella platycephala

o Imantodes splendidus oliveri

o Micrurus nuchalis nuchalis

o Tantilla rubra
Tecpan

u Thorius minutissimus (Santo Tomas)

340 The University Science Bulletin

Tehuantepec (city, and environs)
a Bufo lateralis
a Cystignathus jragilis
a Diaglena reticulata (Cerro Arenal)
a Eleutherodactylus avocalis (Tres Cruces)
a Rhinophrynus rostraius
s Ameiva undulata undulata (by restriction)
s Cnemidophorus deppii deppii (by restriction)
s Cnemidophorus guttatus immutabilis (by present restriction)
B Cnemidophorus guttatus striatus (by present restriction)
s CneniidopJiorus microlepidopus (by present restriction)
s Enyaliosaurus quitiquecarinatus (by restriction)
6 Lepidophyma sinithi tehuanae (Cerro Arenal, 30 kilometers

west of Tehuantepec)
s Phrynosoma spinimentum (by present restriction)
s Sceloporus humeralis (by present restriction)
s Sceloporus melanorhinus melanorhinus (by present restric-
tion)
s Sceloporus siniferus siniferus (by present restriction)
s Sceloporus variabilis smitM (Guengola Mt., 6 kilometers

northwest of)
s Sphaerodactylus inornatus (by present restriction)
s Urosaurus bicarinatus anonymorphus
o Bothrops dunni

o Coniophanes imperialis copei (between Cerro Guengola and)
o Conophis vittatus viduus
o Dipsas elegans

o Dryadophis melanolomus tehuanae (Cerro Guengola)
o Geagras longicaudatVrS (by present restriction)
o Geagras redimitus
o Geophis dubius
o Geophis isthmicus

o Leptodeira mystacina (by present restriction)
o Leptotyphlops phenops phenops
o Loxocemus sumichrasti

o Masticophis mentovarius mentovarius (by present restric-
tion)
o Micrurus ephippifer (by present restriction)
o Oxybelis aeneus auratus (by present restriction)
o Thalerophis diplotropis diplotropis
o Thamnophis ruthveni (3 miles northwest of)
o Trimorphodon biscutatus biscutatus (by present restriction)
o Trimorphodon major
o Tropidodipsas niacdougalli
Tlapancingo

s Gerrhonotus liocephalus liocephalus (by present restriction)
s Gerrhonotus tessellalus (by present restriction)
Totolapam

s Phyllodactylus muralis

Mexican Reptiles and Amphibians 341

o Stenorhina degenhardti qidnquelineata (by present restriction)

o Stenorrhina freminvillii freminvillii (by present resrtiction)
Totontepec

o Geophis anocularis
Ventosa, Bay

t Caretta remivaga

s Gymnophthalmus sumichrasti (by present restriction)
Xuimygopk, between Ayutla and Cacalotepec, 8,650 ft.

o Toluca amphisticha
Zempoaltepec, Mt.

s Abronia fuscolabialis

s Sceloporus cochranae

s Sceloporus siniferus cupreus (by present restriction)

PUEBL.\

Alchichica, Lake

u Ambystoma subsalsum
Cacaloapam

o Bothrops melanurus (by present restriction)

o Trimeresurus garciai
Matamoros (Izucar)

a Engystoma mexicanum (by present restriction)

a Hyla microtis

a Hylodes berkenbuschii (by present restriction)

a Tomodactylus nitidus

s Cnemidophorus meocicanus (by present restriction)

s Phrynosoma taurus

o Leptodeira splendida

o Micrurus laticollaris

o Tachymenis melanocephala (by present restriction)

o Trimorphodon latifascia (by present restriction)
Necaxa

o Lampropeltis triangulum arcijera (by present restriction =
L. doliata arcifera)

o Tantilla morgani
Puebla

a Hyla cardenasi

s Sceloporus spinosus spinosus (by present restriction)

o Chionactis diasii
Rio Frio, 2 miles east of

u Pseudoeurycea bellii (by present restriction)

u Pseudoeurycea cephalica cephalica (by present restriction)
Santa Caterina

s Sceloporus picius
Sierra de Zacapoaxtla

s Gerrhonotus deppii digueti
Tehuacan

a Eleutherodactylus cactorum (20 miles northwest at kilometer
226)

o Salvadora intermedia richardi (1 mile north of)

o Trimorphodon forbesi (San Diego)

342 The University Science Bulletin

Teziutlan

o Rhadinaea quinquelineata

QUERETARO

Jalpan

s Lepidophyma smithi occulor
San Juan del Rio

u Bathysiredon dumerilii queretarensis

QUINTANA ROO

Catmis

o Thamnophis arabdottis
Coba

o Thalerophis mexicanus yucatanensis
Cozumel Island

t Kinosternon cruentatum censors

s Anolis cozumelae

s AristeUiger iuegularis

s Cnemidophorus deppii cozumelus

s Sceloporus cozumelae

o Eutaenia rutiloris

o Thamnophis sauritus chalceus (^ T. sirtalis chalceus, by
present restriction)

San Luis Potosi

No definite locality

o Leptotyphlops humilis tenuiculus
Alvarez

u Chiropterotriton multidentata

s Sceloporus pilsbryi

o Crotalus triseriatus triseriatus (by present restriction)

o Rhadinaea gaigeae
Arroyo Sacahuite (at Palictla, 6 miles north of Tamazunchale)

a Rana moorei
Charcas

s Sceloporus goldmani
Chijol

t Terrapene goldmani
Ciudad Maiz

o Oxybelis potosiensis (38 kilometers northwest of)
Matehuala

s Sceloporus parvus scutulatus (30 miles north of)
San Luis Potosi

o Conopsis nasus heliae

o Geophis latifrontalis (50 miles south of)

o Lampropeltis mexicana

o Tantilla deviatrix (= T. bocourti deviatrix)
Xilitla (vicinity of)

s Gerrhonotus liocephalus loweryi

s Xenosaurus newmanorum

o Adelphicos newmanorum (region)

Mexican Reptiles and Amphibians 343

o Rhadinaea marcellae

o Schmidtophis ruhriventris

o Tantilla shaivi

SiNALOA

Mazatlan

a Bufo kelloggi (2 miles east of)

a Bufo mazatlanensis (2 miles east of)

a Microhyla mazatlanensis (2 miles east of)

t Kinosternon hirtipes (by present restriction)

t Pseudemys scripta ornata

s Anolis nebulosus (by present restriction)

s Anolis utowanae (10 miles north of)

s Callisaurus draconoides bogerti (Los Chivos Island in port
of)

s Ctenosaura teres brachylopha

s Holbrookia maculata elegans

o Lepfodeira personata

o Leptodeira punctata (by present restriction)

o Leptognathus albocinctus (by present restriction)

o Sphenocalamus lineolatus

o Tantilla bimaculata

o Triniorphodon paucimaculata

0 Tropidodipsas philippii

o Tropidonotus quadriserialis
Plomosas

s Sceloporus nelsoni

o Crotalus stejnegeri
Presidio de Mazatlan

a Diaglena spatulata

a Hypopachus oxyrrhinus

a Pternohyla fodiens

a Rana forreri

t Geoemyda pulcherrima pulcherrima (by present restriction)

s Eumeces humilis

s Sceloporus clarkii boulengeri

s Vta lateralis (by restriction)

o Coluber striolatus (by present restriction)

o Gyalopion quadrangularis

o Masticopliis flagellum lineatus (by present restriction)
Rosario

0 Drymarchon corals rubldus
San Bias

o Salvadora hexalepis celeris

Son OR A

No definite locality

s Phrynosoma ditmarsi (not far from Arizona boundary)
Mainland localities
Alamos

s Eumeces parviauriculatus

344 The University Science Bulletin

o Leptodeira ephippiata (Agua Marin, 8.3 miles west-northwest
of)

o Phyllorhynchus hrowni fortitus
El Tigre Mountains

s Sceloporus undulatus virgatus (Santa Maria Mine, in)
Guaymas

t Sphargis coriacea schlegeKi (by present restriction)

s Cnemidophorus burti (LaPosa, 10 miles northwest of)

s Holbrookia maculata thermophila

s Uta gularis

s Uta taylori (10 miles northwest of)

o Caudisona atrox sonoraeiisis (by present restriction)

o Chilomeniscus cinctus

o Gyalopion desertorum (12 kilometers northwest of)

o Masticophis bilineatus (by present restriction)

o Micruroides euryxanthus (by present restriction)

o Tantilla hobartsmithi (10 miles northwest of)

o Trimorphodon lambda
Guirocoba, 18 miles southeast of Alamos, 1,485 ft.

t Pseudemys scripta hiltoni

t Terrapene klauberi

s Callisaurus draconoides brevipes

o Pseudoficimia hiltoni
Hermosillo

s Cnemidophorus tigris aethiops

s Coleonyx variegatus sonoriensis (5 miles southeast of)

s Dipsosaurus dorsalis sonoriensis

s Phyllodactylus homolepidurus (5 miles southwest of)
Llano (between Nogales and Hermosillo)

t Kinosternon flavescens stejnegeri
Los Nogales

s Phrynosoma bufoniiim (by present restriction)

s Urosaurus ornatus linearis
Puerto Libertad

s Phrynosoma platyrhinos goodei (by present restriction)
Santa Magdalena

s Urosaurus ornatus schottii

o Chionactis occipitalis palarostris (5 miles south of)

o Diadophis regalis regalis (by present restriction)
Tepoca Bay

s Uma notata cowlesi (shores of)
Zuiii

s Phrynosoma regale (Sierra de la Nariz, near)
San Esteban

s Cnemidophorus estebanensis

s Ctenosaura conspicuosa

s Sauromalus varius
San Pedro Martir

s Cnemidophorus tigris martyris

s Uta palmeri

Mexican Reptiles and Amphibians 345

San Pedro Nolasco

s Cnemidophorus hacatus

s Uta nolascensis
Tiburon

s Callisaurus draconoides inusitatus

s Cnemidophorus disparilis

s Cnemidophorus punctilineatus

s Crotaphytus dickersonae

s Sauromalus obesus townsendi

Tabasco

No definite localitj'^

t Cinosternum berendtianum

t Claudius agassizii (by present restriction)

t Claudius angustatus

t Claudius megalocephalus (by present restriction)

s Laemanctus deborrei

o Dipsas maxillaris
San Juan Bautista

o Scolecophis scytalinus
Teapa

s Sceloporus teapensis

o Bothrops nummifer nummifer
Tenosique

o Pliocercus elapoides laticollaris (=z P. laticollaris)

Tamaulipas

No definite locality

a Bufo debilis (lower part of the Rio Grande del Norte)

s Holbrookia maculata approximans (lower Rio Grande)
Antiguo Morelos

o Micrurus fitzingeri microgalbineus (7 kilometers south of)
La Clementina, Hacienda, near Forlon

o Hypsiglena dunklei
Marmolejo

s Eumeces dicei
Matamoros

u Diemictylus meridionalis (by restriction)

a Scaphiopus couchii (by restriction)

s Eumeces tetragrammus (by restriction)

s Eumeces tetragrammus funebrosus

o Coniophanes imperialis imperialis

o Lampropeltis triangulum annulata (= L. doliata annulata)

o Zamenis conirostris
Mier

o Hypsiglena ochrorhynchus texana (by present restriction)
Miquihuana

a Eleutherodactylus batrachylus

o Lampropeltis thayeri

346 The University Science Bulletin

Tampico

u Diemictylus kallerti

t Terrapene mexicana mexicana (by restriction)

s Ctenosaura acanthura (by restriction)

s Cyclura shaimi (by restriction)

s Cyclura teres

s Iguana (Ctenosaura) armata (by present restriction)

s Iguana (Ctenosaura) bellii (by present restriction)

s Iguana (Ctenosaura) lanceolata (by present restriction)

o Glaphyrophis lateralis

o Leptotyphlops myopicus myopicus (Savineto)

o Natrix rhombifera blanchardi

c Crocodilus biscutatiis (by present restriction)

c Crocodilus mexicanus
Victoria, Ciudad

s Ameiva undulata podarga (7 miles west of)

o Pliocercus elapoides celatus
Villagran

o Agkistrodon bilineatus taylori (21 kilometers north west of)

Tlaxcala

Huamantla

o Crotalus scutulatus salvini

Veracruz

No exact locality

s Anolis cymbops

o Rhabdosoma guttulatum
Acultzingo

u Thorius dubitus (2 miles west of)

u Thorius troglodytes (2 miles west of)

a Hyla arborescandens (3 miles west of)

a Hyla bistincta (Lake San Bernardino, by present restriction)

a Hyla forbesi (3 miles west of)

s Sceloporus formosus formosus (by present restriction)

s Sceloporus mucronatus aureolus (2 miles west of)

o Geophis blanchardi (2 miles west of)

o Toluca lineata varians (by present restriction)
Alvarado

t Dermatemys mawii (by present restriction)

t Emys callirostris (by present restriction)

t Limnochelone micrura (by present restriction)

t Pseudeniys scripta cataspila (by present restriction)

t Staurotypus triporcatus
Atoyac

a Eleutherodactylus alfredi

o Phrynonax guentheri
Cabo Rojo, near Panaco Island

o Crotalus durissus totonacus
Cascajal, Upper Uzpanapa River

s Anolis barkeri

Mexican Reptiles and Amphibians 347

Castillo deTeayo

o Coniophanes fissidens convergens (6 miles northwest of)
Cerro Gordo

s Gerrhonotus lemniscatus (by present restriction)
Cordoba

a Agalychnis callidryas

a Hyla euphorbiacea (by present restriction)

a Hylodes sallaei (by present restriction)

a Microbatrachylus albolabris (2 miles west of)

a Microhyla elegans

a Smilisca baudinii baudinii (by present restriction)

a Smilisca daulinia (by present restriction)

s Iguana iguana rhinolopha (by present restriction)

s Leiolopisma silvicolum (San Lorenzo, 10 miles southeast of)

s Xenosaurus grandis

o Boa diviniloquax mexicana (by present restriction)

o Constrictor constrictor imperator (by present restriction)

o Drymobius margaritiferus margaritiferus (by present re-
striction)

o Geophis semidoliatus (by present restriction)

o Micrurus affinis afRnis (by present restriction)

o Pseustes poecilonotus argus (by present restriction)

o Stenorrhina degenhardtii mexicana (by present restriction)

u Thamnophis phenax phenax

o Typhlops praelongus
Cosamaloapam

a Acrodytes spilomma

t Cinostemon effeldtii (by present restriction)

t Kinosternon acutum (by present restriction)

t Kinosternon leucostomum (by present restriction)

t Swanka maculata (by present restriction)
Cruz Blanca

u Spelerpes laticeps (by present restriction)

s Sceloporus aeneus bicanthalis

s Sceloporus dispar (by present restriction)
Cuatotolapam

g Dermophis mexicanus mexicanus (by present restriction)
Cuautlapan

u Pseudoeurycea nigromaculata

a Eleutherodactylus natator

a Eleutherodactylus spatulatus

a Hyla dendroscarta

o Chersodromus liebmanni (by present restriction)

o Chersodromus nigricans (by present restriction)

o Dirosema collars (by present restriction)

o Rhadinella schistosa

o Tantilla phrenitica

o Scaphiodontophis cyclurus
Encero

s Sceloporus serrifer plioporus (4 miles east of)

348 The University Science Bulletin

Huatusco

a Syrrhophus verruculatus

s Anolis lemurinus bourgaei (by present restriction)

s Anolis tropidonotus

s Xenosaurus fasciatus

o H omalo cranium boulengeri

o Opisthiodon torquatus
Jalapa

u Bolitoglossa platydactyla (by present restriction)

u Parvimolge townsendi (Cerro de los Estropajos, near)

u Pseudoeurycea gigantea

a Bufo cristatus

a Bvjo occipitalis (by present restriction)

a Eleutherodactylus decoratus (Banderilla, 6 miles west of)

a Eleutherodactylus dunni (Cerro de los Estropajos, near)

a Eleutherodactylus venustus

a Hyla miotympanum (by present restriction)

a Hyla nigropunctata

a Hyla picta

a Hyla taeniopus

a Hylodes plicatus

a Syrrhophus mystaceus (Cerro de los Estropajos, near)

s Anelytropsis papillosus

s Anolis laeviventris (by present restriction)

s Anolis sallaei (by present restriction)

s Anolis sericeus (El Encero de)

s Anolis wiegmanni (by present restriction)

s Celestus enneagrammus

s Corylhophones chamaeleopsis (by present restriction)

s Corythophanes hernandezii (by present restriction)

s Corythophanes mexicanus (by present restriction)

s Eumeces lynxe furcirostris

s Laemanctus longipes

s Sceloporus jalapae

s Sceloporus malachiticus salvini (by restriction)

o Coniophanes fissidens proterops

o Crotalus durissus durissus (by present restriction)

o Elapochrous deppei (by present restriction)

o Gcophis fuscus

o Lampropeltis triangulum polyzona (Cuatupe, near)
(= L. doliata polyzona)

0 Leptodeira frenata

o Leptodeira maculata (5 miles east of, by present restriction)

o Liophis tricinctus (by present restriction)

o Micrurus elegans elegans (by present restriction)

o Pituophis deppei lineaticollis

o Pliocercus elapoides elapoides

o Thamnophis scalaris scalaris
Jicaltepec

o Adelphicos quadrivirgatus quadrivirgatus (by present re-
striction)

Mexican Reptiles and Amphibians 349

o Adelphicos quadrivirgatus aculirostrurn (by present restriction)

o Coluber sihon (by present restriction)

o Sibon nebulatus (by present restriction)

o Storeria dekayi temporalineata (San Rafael)
Jico (Xico)

a Hyla probosddea (2 miles west of) (^ Hyla dalquesti)
La Joya

u Chiropterotriton lavae (2 miles west of)
La Laja

t Kinosternon herrerai
Metlac

u Oedipina lineola (by present restriction)

u Spelerpes infuscatus (by present restriction)
Mirador

u Chiropterotriton chiroptera

a Hyla gracilipes

a Hyla muricolor

s Sceloporus heterurus

s Sceloporus mucronatus mucronatus

s Sceloporus viviparus

o Bothrops nummifer veraecrucis

o Coluber novae hispaniae (by present restriction)

o Galedon annularis (by present restriction)

o Imantodes cenchoa leucomelas

o Leptognathus dumerili (by present restriction)

o Spilotes pullatus auribundus

o Spilotes pullatus mexicanus (by present restriction)

o Spilotes variabilis (by present restriction)

o Tantilla miniata

o Tropidodipsas sartorii sartorii
Misantla

a Hyla godmani (by present restriction)
Orizaba

u Bolitoglossa rufescens

u Pseudoeurycea leprosa

u Spelerpes gibbicaudus

u Spelerpes orizabensis (Mount)

u Thorius pennatulus

a Eleutherodactylus beatae (La Perla, near)

a Eleutherodactylus rhodopis (by restriction)

a Hyla staufferi

s Abronia taeniata graminea

s Barisia antauges

s Barisia modesta (by present restriction)

s Corythophanes cristatus (by present restriction)

s Diploglossus chalybeus

s Gerrhonotus liocephalus ophiurus

s Laemanctus serratus (Valley of)

s Leiolopisma gemmingeri

s Phrynosoma orbiculare cortezii (Hacienda del Jasmin, be-
tween Cordoba and)

350 The University Science Bulletin

o Bothrops undulatus (by present restriction)

o Drymarchon corais orizabensis

o Flops corallinus crebripunctatus (by present restriction) *

o Ficimia olivacea (by present restriction)

o Leptodeira annulata taylori

o Ninia diademata diademata

o Rhadinaea fulvivittis (Alpine region of)

o Rhadinaea vittata (by present restriction)

o Storeria dekayi anomala

o Streptophorus bijasciatus (by present restriction)

o Thamnophis chrysocephalus

o Thamnophis sumichrasti sumichrasti

o Typhlops basimaculatus (by present restriction)

o Typhlops perdiius
Pan de 011a (south of Tezuitlan)

a Hyla pachyderma

o Toluca lineata wetmorei
Perez

s Cnemidophorus deppii oligoporus
Potrero Viejo

a Hyla rickardsi

a Leptodactylus labialis (by present restriction)

a Syrrhophus cystignathoides

a Eumeces sumichrasti (by restriction)

s Leiolopisma cherriei stuarti

o Coniophanes imperialis clavatus (by present restriction)

o Oxyrhopus baileyi

o Rhadinaea decorata (by present restriction)

0 ScapJuodontophis nothus

o Thaleropliis mexicanus mexicanus (by present restriction) ♦
Puerto Mexico

a Bujo eiteli

o Calopisma quinquevittatum mexicana (by present restriction)

o Hydrops lubrictis (by present restriction)
Rodriguez Clara

a Microbatrachylus pygmaeus

o Liophis varia (by present restriction)
San Andres Tuxtla

s Ameiva undulata amphigramma

o Coniophanes Rssidens Rssidens (by present restriction)
San Jose Acateno

o Geophis longiceps
San Lorenzo

o Ficimia publia taylori
Tecolutla

s Holbrookia propinqua piperata (2 miles south of)

• No specimens are definitely known from the state of Puebla.

* Leptophis mexicanus mexicanus Smith and Taylor, Bull. U. S. Nat. Mus., no. 187, 1945,
p. 91.

Mexican Reptiles and Amphibians 351

Tequejoitepec (7 miles west of Jalapa)
a Eleutherodactylus dorsoconcolor
o Rhadinaea forbesi

Totalco

u Pseudoeurycea melanomolga (20 kilometers north of)

s Sceloporus niegalepidurus

o Crotalus gloydi lautus

o Sistrurus ravus (by present restriction)

o Thamnophis eques eques (by present restriction)
Tuxpan

o Pliocercus bicolor
Tuxpango (near Orizaba)

o Trimorphodon collaris
Veracruz

a Bufo marmoreus

a Bufo horribilis

a Bujo trachypus (by present restriction)

a Bufo valliceps (by present restriction)

a Rana pipiens austricola (by restriction)

a Rhinophrynus dorsalis

t Emys berardii

s Anolis jacobi (by present restriction)

s Basiliscus vittatus (by present restriction)

s Cnemidophorus guttatus guttatus Wiegmann (by present re-
striction)

s Ctenosaura cycluroides (by present restriction)

s Cyclura denticulata (by present restriction)

s Cyclura semicristata (by present restriction)

s Sceloporus variabilis variabilis (by present restriction)

o Conophis lineal us lineatus (by present restriction)

o Ninia sebae sebae (by present restriction)

o Streptophorus sebae collaris (by present restriction)

c Crocodilus americanus (by present restriction)
Xometla (8,500 ft., Mt. Orizaba)

u Pseudoeurycea gadovii
Zacuapan

o Dryadophis melanolomus veraecrucis

o Eudryas boddaerti mexicanus

Yucatan

No specific locality

u BoUtoglossa yucatana

a Eleutherodactylus laticeps
Chichen Itza

a Triprion petasatus (by present restriction)

t Kinosternon creaseri (1 mile south of Hacienda)

t Terrapene mexicana yucatana (by present restriction)

s Anolis aureolus (by present restriction)

s Anolis beckeri (by present restriction)

s Cnemidophorus sackii angusticeps (by present restriction)

352 The University Science Bulletin

s Enyaliosaurus defensor (by present restriction)

s Leiolopisma cherriei ixbaac

s Norops yucatanicus

s Sceloporus chrysostictus (by present restriction)

s Thecadactylus rapicaudus (by present restriction)

o Bothrops yucatanicus

o Cochliophagus tomieri (by present restriction)

o Coniophanes schmidti

o Conophis Uneatus concolor (by present restriction)

o Dipsadomorus fasciatus (by present restriction)

o Dipsas hrevifacies (by present restriction)

o Dipsas sanniolus (by present restriction)

o Dryadophis melanolomus melanolomus (by present restric-
tion)

o Drymarchon corais melanocercus (by present restriction)

o Drymarchon corais melanurus (by present restriction)

o Elaphe flavirufa flavirufa (by present restriction)

o Ficimia ornata (by present restriction)

o Ficimia publia (by present restriction)

o Geophis multitorque.s yucatanicus

o Imantodes splendidus splendidus (by present restriction)

o Imantodes tenuissimus (by present restriction)

o LampropeUls triangulum bhmchardi (= L. doliata blanchardi)

o Leptodeira yucatanensis yucatanensis (by present restric-
tion)

o Leptognathus leucostomus (by present restriction)

o Leptognathus subannulatus (by present restriction) '

o Leptognathus torquatus (by present restriction)

o Micrurus affinis mayensis

o Ninia sebae morleyi

o Oxybelis fulgidus (by present restriction)

o Pliocercus elapoides schmidti

o Tantilla canula (by present restriction)

o Tropidodipsas fasciata (by present restriction)

o Typhlops niicrostomus (by present restriction)
Citilpech

s Anolis acutirostris
Dzitas

o Opheodrys mayae
Libre Union

o Pliocercus andrewsi
Merida

s Anolis kidderi

s Laemanctus alticoronatus

s Sceloporus lundelli gaigeae

s Sceloporus serrifer serrifer (by present restriction)

s Sphaerodactylus glaucus glaucus

o Coniophanes meridanus

o Tantilla cuniculator

Mexican Reptiles and Amphibians 353

Progreso

s Ameiva undulata gaigeae

s Hemidactyhis exsul

Zacatecas

Hacienda El Florencio

a Eleutherodactylus occidentalis
La Colorada

o Crotalus molossus nigrescens (4 miles west of)
Valparaiso Mts.

s Sceloporus jarrovii minor (by present restriction)

UNITED STATES

Arizona

Apache County

Springerville, 11 miles south of
a Hyla wrightorum
Cochise County
Chiricahua Mts.

s Urosaurus ornatus chiricahuae (Pinery Canyon)
Fairbank

s Cnemidophorus arizotiae
Huachuca Mts.

s Holbrookia maculata pulchra (Carr Canyon, 5,200 ft.)
s Sceloporus jarrovii jarrovii (by present restriction)
s Sceloporus scalaris slevini (Miller Peak)
o Crotalus lepidus klauberi (Carr Canyon)
o Crotalus triseriatus pricei (= C. pricei pricei)
o Crotalus willardi (Ramsey Canyon; by present restric-
tion)
o Tantilla wilcoxi wilcoxi (Fort Huachuca)
Coconino County

Painted Desert, Little Colorado River
s Crotaphytus collaris baileyi
Gila County

Tonto Creek, 6,000 ft.

o Thamnophis rufipunctatus
Graham County
Camp Grant

o Bascanium semilineatus (by present restriction)
o Masticophis flagellum piceus
Gila Mts.

s Sauromalus obesus tumidus (Telegraph)
Maricopa County
Wickenburg

o Crotalus scutulatus scutulatus (by present restriction)
Mohave County

Fort Mojave, bet-ween Fort Yuma and, along the Colorado River
near 35° N.
a Bufo frontosus
22—90

354 The University Science Bulletin

Navajo County

Winslow, 10 miles east of

o Arizona elegans philipi
Pima County

Fort Buchanan (4 miles south of Tucson)

o Gyalopion canum

o Hypsiglena chlorophaea
Indian Oasis, 27 miles west of

o Lampropeltis getulus yumensis
Santa Catalina Mts.

o Diadophis regalis arizonae (Sabino Canyon)
Sasabe

o Crotalus tigris
Sierra cle Morena (= Sierra de la Union, 10 miles southwest of
San Miguel)

s Heloderma suspectum
Tucson

t Cinosternum punctatuni (by present restriction)

t Kinosternon sonoriense

s Callisaurus draconoides ventralis (by present restriction)

o Diadophis regalis laetus (by restriction)

o Phyllorhynchus broivni browni

o Pituophis catenifer ruiilis

o Thamnophis macrostemma megalops (= T. subcarinatus
megalops) (by present restriction)
Xavier

s Coleonyx variegatus bogerti

o Phyllorhynchus decurtatus nubilis (Weisner's Ranch)
Santa Cruz County

Calabasas Canyon (4 miles north of Mexican border, south of
Calabasas)

o Oxybelis microphthalmus
Santa Rita Mts *

a Hyla affinis

a Hyla arenicolor (by present restriction)

s Sceloporus clarki clarki (by present restriction)

o Lampropeltis getulus splendida (by present restriction)

o Salvadora grahamiae (by present restriction)

o Sonora semiannulata semiannulata (by restriction)
Yavapai County
Canyon Prieto

o Crotalus mitchellii pyrrhus
Fort "Whipple

o Lampropeltis pyromelana

0 Salvadora hexalepis hexalepis

o Sonora semiannulata isozona

• While the type localities of all forms listed for this locality were originally stated as
"Sonora," with little doubt the specimens actually were secured in the part of Arizona that
was only long ago part of Sonora. Stickel (Proc. Biol. Soc. Wash., vol. 56, 1943, p. 120) pro-
vides reasonable evidence for justification of his restriction of the type locality of Sonora s.
semiannulata to the vicinity of the Santa Rita Mountains. We believe it equally reasonable to
so restrict the other names here cited.

Mexican Reptiles and Amphibians 355

Yuma County
Yuma

a Bufo alvarius (by present restriction)

s Cnemidophorus gracilis (by present restriction)

s Cnemidophorus melanostethus (by present restriction)

s Phrynosoma solare (by present restriction)

s Sceloporus magister magister (Fort Yuma)

s Uma notata notata (by present restriction)

s Uma rufopunctata

Arkansas

No specific locality

o Elaphe laeta laeta (Red River)
o Thamnophis marciana (Red River)
Lawrence County
Imboden

u Siren intermedia nettingi

California

No specific locality

s Crotaphytus gambelii

s Eumeces skiltonianm amhlygrammus (Fort Humboldt)
o Masticophis lateralis
El Dorado County
No specific locality

o Lampropeltis getulus boylii
Fresno County
Fort Miller

s Cnemidophorus tigris mundus
s Cnemidophorus undulatus
Fresno

o Tantilla eiseni eiseni
Imperial County
Holtville

o Chionactis occipitalis annulatus (by present restriction)
Seeley

o Sonera semiannulata linearis
Winterhaven (= Fort Yuma, Camp Yuma)

s Coleonyx variegatus variegatus

s Dipsosaurus dorsalis dorsalis

s Phrynosoma m'callii (between Vallecita and)

s Sauromalus obesus obesus

s Urosaurus graciosus

s Urosaurus ornatus symmetricus
Inyo County
Death Valley

s Anota calidiarum
Kern County
El Paso Creek

s Sceloporus occidentalis biseriatus (borders of, by present
restriction)

356 The University Science Bulletin

Fort Mohave, Mountains near

t Gopherus agassizii
Fort Tejon

u Ensatina croceater

s Xantusia vigilis
Randsburg

o Lichanura roseofusca gracia
Tehachapi Mountains

s Eumeces gilberti rubricaudata
Los Angeles County

Pasadena, Arroyo Seco Canyon

s Uta stansburiana hesperis
Monterey County
Monterey

u Aneides lugubris lugubris
Orange County

Lower Coyote Creek, near Alamitos

t Clemmys marmorata pallida
San Benito County
Bear Valley

s Phrynosoma coronatum frontale
San Bernardino County
Mojave River

s Eumeces quadrilineatus (by present restriction)
San Diego County
Boulder Creek

u Taricha klauberi
Coast Range, summit of

s Sceloporus graciosus vandenburgianus

s Streptosaurus mearnsi (at boundary of United States and
Mexico)
Deerhorn Flat

o Salvadora hexalepis virgultea
Dry Lake, Bensons

o Arizona elegans eburnata

o Phyllorhynchus decurtatus perkinsi
Dulzura

o Crotalus ruber (by present restriction)
La Jolla

o Arizona elegans occidentalis
Milquatay Valley

s Sceloporus orcutti orcutti
Mountain Spring, Colorado Desert

o Bascanion flagellum frenatum
Narrows, The

o Crotalus cerastes laterorepens

o Rhinocheilus antonii clarus (Borego Valley, 2 miles north
of)
Poway

s Xantusia picta (by present restriction)

Mexican Reptiles and Amphibians 357

i'roctor Valley, between Jamul and Upper Otay Reservoir

s Coleonyx variegatus abbotti
San Diego

s Anniella pulchra pulchra (by present restriction)

s Anniella texana (by present restriction)

s Elgaria multicarinata webbii (by restriction)

s Phrynosoma coronatum blainvillii (by present restriction)

o Coronella multifasciata (by present restriction)

o Diadophis amabilis similis

o Laynpropeltis calijomiae (= Lampropeltis getulus cali-
forniae) (by present restriction)

o Pituophis catenifer annectens

o Rhinocheilus lecontei lecontei

o TImmnophis harmnondii (= T. elegans hammondii) (by
restriction)
Vallecito, 35 miles west of

o Leptotyphlops humilis humilis (by present restriction)
Wildwood Ranch, 1,520 ft. (5 miles southwest of Ramona)

o Trimorphodon vandenburghi
Witch Creek, 2,700 ft.

s Xantusia henshawi
Yaqui Well

o Leptotyphlops humilis cahuilae

o Tantilla eiseni transmontanus (1 mile east of)
San Francisco County
San Francisco

a Rana aurora draytoni (by present restriction)
Santa Barbara County
Santa Barbara

o Lampropeltis zonata zonata
Shasta County

Fort Reading (near Redding)

a Scaphiopus hammondii
Solano County
Benicia

a Bufo boreas halophilus
Ventura County

Santa Paula, 800 ft.

a Bufo calif ornicus

COLOKADO

Fremont County
Beaver Creek

s Cnemidophorus tesselatus
Larimer County
Cow Creek

s Eumeces multivirgatus
Prowers County
Arkansas River

a Bufo cognatus

358 The University Science Bulletin

Pueblo County
Pueblo

o Masticophis ttagellum testaceus (by present restriction)

Floeida

Monroe County
Key West

t Lepidochelys kempii (by present restriction)

Indiana

Posey County

New Harmony, Fox River at

t Pseudemys scripta elegans

Iowa

Pottawatomie County
Council Bluffs

t Terrapene ornata (by present restriction)

Kansas

Cowley County
Winfield

o Sistrurus catenatus tergeminus (by present restriction)
Geary County
Fort Riley

a Microhyla olivacea (by present restriction)
s Phrynosoma cornutum (by present restriction)
s Phrynosoma harlanii (by present restriction)
Riley County
Manhattan

t Chrysemys picta bellii (by present restriction)

Louisiana

Orleans Parish
New Orleans

t Chelydra serpentina (by present restriction)

Missouri

Cape Girardeau County

Cape Girardeau, below, on Mississippi River
s Leiolopisma laterale
Jasper County
Carthage

o Pituophis catenifer sayi (by present restriction)

Montana

Chouteau County
Fort Benton

a Bufo dipternus (by present restriction)

Nebraska

Boyd County
Gross

o Crotalus viridis viridis (by present restriction)

Mexican Reptiles and Amphibians 359

Washington County

Boyers Eiver, stone quarry on west side of Missouri River three
miles above the mouth of
o Thamnophis sauntus proximus (= T. sir talis proximus)
o Thamnophis sirtalis parietalis {= T. ordinatus parietalis)

New Jersey

Gloucester County
Raccoon

a Rana pipiens

New Mexico

No definite locality

u Siredon harlanii (Spring Lake)
t Cinosterniim henrici
Dona Ana County
Jornada del Muerto

s Crotaphytus jasciatus
s Laniprosaurus guttulatus
Valley, 20 miles northeast of Fort Cummings

s Holbrookia maculata flavilenta
Las Cruces

s Phrynosoma modestum (by present restriction)
Grant County
Fort Webster

s Elgaria kingii nobilis (Copper mines of the Gila)
o Crotalus molossus molossus (Santa Rita del Cobre)
McKinley County
Zuiii

o Pituophis catenifer affinis
Otero County
Alamogordo

s Uta stansburiana stejnegeri (mouth of dry canyon)
San Juan County
Pueblo Bonito

s Cnemidophorus gularis velox (by present restriction)
Shiprock

o Masticophis taeniatus taeniatus (by present restriction)
Santa Fe County
Santa Fe

s Gambelia wisUzenii wislizenii

a Phrynosoma douglassii hernandezi (by present restric-
tion)
Sierra County
Lake Valley

o Leptotyphlops myopicus dissectus

New York

Albany County
Albany

a Acris crepitans (by present restriction)

360 The University Science Bulletin

Oklahoma

Beckham County
Suydam Creek

s Sceloporus undulatus consobrinus (junction with nr.rii,
fork of Red River)
Garvin County
Maysville

o Psammophis flavigularis (by present restriction)
Tulsa County
Tulsa

o Natrix erythrogaster transversa (by present restriction)
o Natrix rhombifera rhambifera (by present restriction)
Oregon

Wasco County
The Dalles

s Eumeces skiltonianus (by present restriction)
o Crotalus viridis oreganus (by present restriction)

South Carolina

Charleston County
Charleston

a Rana catesbeiana (by present restriction)

t Testudo cephalo (by present restriction)

t Testudo viridis (by present restriction)

s Cnemidophorus sexlineatus (by present restriction)

o Opheodrys aestivus (vicinity of; by present restriction)

Tennessee

Hardeman County

Boliver, 10 miles northeast of

o Agkistrodon piscivorus leucostomus (by restriction)
Tejxas

Aransas County
Rockport

t Malaclemys terrapin littoralis
Atascosa County

Pleasanton, 9 miles east of
0 Tanlilla kirnia

o Tantilla nigriceps fumiceps (by restriction)
Somerset, 9 miles southwest of

s Holbrookia propinqua (by present restriction)
Bexar County
Helotes

a Eleutherodactylus latrans
s Coleonyx brevis
s Eumeces brevilineatus
San Antonio

u Ambystoma tigrinum proserpine (Salado River, 4 miles

east of)
a Bufo nebulifer (by present restriction)

Mexican Reptiles and Amphibians 361

t Pseudemys floridana texana

s Sceloporus variabilis marmoratus

Brewster County

Boquillas, on Rio Grande River

t Pseudemys scripta gaigeae
Chalk Draw

o Leptotyphlops humilis segregus
Chisos Mts.

s Sceloporus merriami annulatus (east slope of)

o Agkistrodon mokeson pictigaster (== A. contortrix picti-

gaster) (Maple Canyon, 5,200 ft.)
o Diadophis regalis blanchardi (the Basin)
o Salvadora hexalepis deserticola (Government Spring)
o Sonora semiannulata blanchardi (northern slopes of)
Calhoun County
Indianola

o Crotalus atrox
Cameron County
Brownsville

a Bufo compactilis speciosus (by present restriction)
a H7jla vanvlietii
a Syrrhophus campi
t Amy da emoryi (Rio Grande River)
t Gopherus berlandieri (by present restriction)
s Cnemidophorus guttatus Hallowell (by present restriction)
s Cnemidophorus sackii gularis (by present restriction)
o Coluber constrictor stejnegerianus (by present restric-
tion)
o Heterodon nasicus kennerlyi (bj^ present restriction)
o Leptodeira annulata septentrionalis (by present restric-
tion)
o Masticophis taeniatus ruthveni
Comal County
New Braunfels

s Holbrookia texana

0 Micrurus fulvius tenere (by present restriction)
Comanche County
Comanche

o Leptotyphlops dulcis (by present restriction)
Duval County
San Diego

a Hypopachus cuneus cuneus
s Cnemidophorus gularis sericeus
s Lysoptychus (Sceloporus) lateralis
o Sonora taylori (by present restriction)
El Paso County
El Paso

a Hyla copii

s Cnemidophorxis marmoratus (by present restriction)

s Phrynosoma planiceps (by present restriction)

362 The University Science Bulletin

Fort Bliss

o Tantilla nigriceps nigriceps (by present restriction/
Erath County-
No definite locality

s Holhrookia maculata lacerata (by present restriction)
Hidalgo County
Hidalgo

s Sceloporus grammicus disparilis (Lomita Ranch, 6 miles
north of)
Jeff Davis County
Davis Mountains

o Elaphe sclerotica
o Elaphe subocularis
o Lampropeltis alterna
Fort Davis

s Cnemidophorus grahami (by present restriction)
s Urosaurus ornatus schmidti
o Coluber taeniatus girardi (by present restriction)
o Elaphe bairdi

o Masticophis taeniatus ornatus (by present restriction)
Kendall County

Edge Falls, 4 miles south of Kendalia
o Storeria dekayi texana
Kleburg County
Kingsville

o Salvadora lineata
McLennan County
Waco

t Kinosternon flavescens Aavescens (by present restriction)
Maverick County
Eagle Pass

o Arizona elegans elegans (by present restriction)

o Coluber arizonae (by present restriction)

o Drymarchon corais erebennus

o Drymarchon corais obsoleta

o Masticophis taeniatus schotti

o Sonora episcopa (by present restriction)

Presidio County
Marfa

s Cnemidophorus septemvittata (by present restriction)
Presidio (del Norte)

o Crotalus lepidus lepidus (by present restriction)
Starr County

Rio Grande City

s Sceloporus olivaceus (Arroyo Los Olmos, 3 miles south-
east of)
o Ficimia streckeri (Arroyo Los Olmos, 3 miles southeast
of)

Mexican Reptiles and Amphibians 363

Val Verde County

Devils River (Rio San. Pedro)

a Bufo punctatus

s Cnemidophorus perplexus perplexus

s Eumeces obsoletus

s Gerrhonotus liocephalus infernalis

s Holbrookia affinis

s Sceloporus poinsettii

s Urosaurus ornatus ornatus (by restriction)
East Painted Cave, near mouth of Pecos River

s Sceloporus merriami merriami
Webb County
Laredo

s Crotaphytus reticulatus (by present restriction)

Utah

Salt Lake County
Great Salt Lake

s Cnemidophorus tigris tigris (valley of)
s Phrynosoma platyrhinos platyrliinos
Washington County

Beaverdam Mountains

o Pituophis catenifer deserticola (by restriction)

Washington
King County
Seattle

a Hyla regilla (by present restriction)

SOUTH AMERICA

ARGENTINA

Buenos Aires

o Bothrops atrox dints

BRAZIL
Amazon River

a Hyla lichenosa (by present restriction)

a Rana palmipes

Espiritu Santo River

o Oxybelis acuminatus
Rio de Janeiro

s Scincus agilis

COLOMBIA
No definite locality

o Bothrops schlegelii

Baranquilla

s Anolis sulcifrons

s Goniodactylus braconnieri
Rio Magdalena

c Caiman crocodilus fuscus

364 The University Science Bulletin

Rio Truando

a Engystomops pustulosus

s Anolis pentaprion
Turbo

a Hyla phaeota

DUTCH GUIANA
No definite locality

o Clelia clelia clelia

ECUADOR

No definite locality

a Bufo intermedius (Andes)
Guayaquil

o Pseudodipsas jallax (by present restriction)

URUGUAY

Montevideo

o Erythrolamprus bizona (by present restriction)

VENEZUELA
No definite locality

a Bujo sternosignatus
Caracas

o Comastes quincunciatus

WEST INDIES

CUBA
Cienfuegos

s Anolis sagrei sagrei (by present restriction)

JAMAICA

No definite locality

t Testudo caouana (by present restriction)

ST. VINCENT
No definite locality

s Hemidactylus mabouia (by present restriction)

s Mabuya mabouya mabouya (by present restriction)

SANTO DOMINGO
Puerto Plata

c Crocodylus acutus acutus (by present restriction)

Mexican Reptiles and Amphibians 365

BIBLIOGRAPHY
Blair, Albert P.

1947. A new Rana from San Luis Potosi, Mexico. Amer. Mus. Novitates
No. 1353, pp. 1-17, figs. 1-8.

BoGERT, Charles M.

1945. Two additional races of the patch-nosed snake, Salvadora hexalepis.
Amer. Mus. Novitates No. 1285, pp. 1-14, figs. 1-10.

• , and James A. Olr'er

1945. A preliminary analysis of the herpetofauna of S'onora. Bull. Amer.
Mus. Nat. Hist., vol. 83, pp. 297-426, figs. 1-13, pis. 30-37, table 1,
maps 1, 2.

Burger, W. Leslie, and M. Max Hensley

1949. Notes on a collection of reptiles and amphibians from northwest-
ern Sonora. Bull. Chicago Acad. Sci., vol. — , pp. — .

, and Wm. B. Robertson

1950. A new .subspecies of the Mexican moccasin Agkistrodon bilineatus.
Univ. Kansas Sci. Bull., vol. 54, 1950 (in press).

CoNANT, Roger

1946. Studies on North American water snakes — II. The subspecies of
Natrix valida. Amer. Midi. Naturalist, vol. 35, pp. 250-275, figs. 1, 2.

Gloyd, Howard K.

1948. Description of a neglected subspecies of rattlesnake from Mexico.
Nat. Hist. Miscellanea, No. 17, pp. 1-4, figs 1-3.

Klauber, L. M.

1946. The glossy snake, Arizona, with descriptions of new subspecies.
Trans. San Diego Soc. Nat. Hist., vol. 10, pp. 311-398, pis. 7-8,
fig 1, map.

1946. The gopher snakes of Baja California, with descriptions of new

subspecies of Pituophis catenifer. Trans. San Diego Soc. Nat. Hist.,
vol. 11, pp. 1-40, pis. 1-2, map.

1947. Classification and ranges of the gopher snakes of the genus Pitu-
ophis in the western United States. Bull. Zool. Soc. San Diego, no.
22, pp. 1-83, figs. 1-6.

Maldonado-Koerdell, Manuel

1948. Los colecciones de anfibios del Museo Alfredo Duges en la Univer-
sidad de Guanajuato I. Urodelos. Mem. Rev. Acad. Nac. Cienc, vol.
56 (2-3) pp. 185, 226, figs. 1-8.

Mertens, Robert

1941. Eine neue Korallennatter aus Mexiko. Senckenbergiana, vol. 25,
pp. 216-217, fig. 1.

Peters, James A.

1948. The northern limit of the range of Lacmanctus serratus. Nat. Hist.
Miscellanea, No. 27, pp. 1-3, fig. 1.

Reese, Robert W., and I. Lester Firschein

1950. Herpetological results of the University of Illinois field expedition,
spring 1949. II. Amphibia. Trans. Kansas Acad. Sci. Bull., vol. 53
(in press).

Schmidt, Karl P., and Frederick A. Shannon

1947. Notes on amphibians and reptiles of Michoacan, Mexico. Fieldiana,
Zool., vol. 3, pp. 63-84, fig. 8.

Shannon, Frederick A., and Hob..\rt M. Smith

1949. Herpetological results of the University of Illinois field expedition,
spring 1949. I. Introduction, Testudines, Serpentes. Trans. Kansas
Acad. Sci., vol. 52 (in press).

366 The University Science Bulletin

Smith, Hob art M.

1946. Preliminary notes and speculation on the triseriatns group of rat-
tlesnakes in Mexico. Univ. Kan. Sci. Bull. vol. 31, pp. 75-101, pi.
3, figs 1-23.

1946. Notas sobre una coleccion de reptiles y anfibios de Chiapas, Mex.
Rev. Soc. Mex. Hist. Nat., vol. 7. pp. 63-74.

1947. Subspecies of the sonoran toad (Bufo compactilis Wiegmann). Her-
petologica, vol. 4, pp. 7-13, fig. 1.

1947. Notes on Mexican amphibians and reptiles. Journ. Wash. Acad.
Sci., vol. 37, pp. 408-412.

— , and Dave A. Langebartel

1950. Notes on a collection of reptiles and amphibians from the Isthmus
of Tehuantepec, Oaxaco. Journ. Washington Acad. Sci., 1950 (in
press) .

, and Leonard E. Laupe

1945. Notes on a herpetological collection from Oaxaca. Herpetologica,

vol. 3, pp. 1-13, fig. 1.
1945. Mexican amphibians and reptiles in Texas Wildlife Collections.

Trans. Kan. Acad. Sci., vol. 48, pp. 325-354.

, C. William Nixon, and Philip W. Smith

1949. Mexican and Central American garter snakes (Thamnophis) in the
British Museum of Natural History. Journ. Linn. Soc. London, vol.
— , pp. — .

, and Edward H. Taylor

1945. An annotated checklist and key to the snakes of Mexico. Bull. U.
S. Nat. Mus. no. 187, pp. i-iv, 1-239.

1948. An annotated checklist and key to the amphibia of Mexico. Bull.
U. S. Nat. Mus., no. 194, pp. i-iv, 1-118.

1950. An annotated checklist and key to the reptiles of Mexico exclusive
of the snakes. Bull. U. S. Nat. Mus., no. 199, pp. i-iv, 1 — (in press).

Smith, Philip W., and W. Leslie Burger

1950. Herpetological results of the University of Illinois field expedition,
spring 1949. III. Sauria. Trans. Kansas Acad. Sci., vol. 53 (in
press).

Tanner, Wilmer W.

1946. A taxonomic study of the genus Hypsiglena. Great Basin Natural-
ist, vol. 5, pp. 25-92, pis. 1-4.

Taylor, Edvv^ard H.

1947. A bibliography of Mexican amphibiology. Univ. Kan. Sci. Bull.,
vol. 31, pp. 543-589.

1948. A new hylid frog from eastern Mexico. Univ. Kan. Publ. Mus. Nat.
Hist., vol. 1, pp. 257-264, 1 fig.

1949. A Preliminary account of the herpetology of the State of San Luis
Potosi, Mexico. Univ. Kansas Sci. Bull., vol. 33, pt. 1, pp. 169-215

1949. New or unusual Mexican amphibians. Amer. Mus. Novitates, no.
1437, Dec. 7, pp. 1-21, figs. 1-6.

1950. Second contribution to the herpetology of San Luis Potosi. Univ.
Kansas Sci. Bull., vol. 33, pt. 2, pp.

Woodbury, Angus M., and Dixon M. Woodbury

1944. Notes on Mexican snakes from Oaxaca. Journ. Wash. Acad. Sci.,
vol. 34, pp. 360-373.

SPECIES INDEX

PAGE

Abronia deppii 332

Abronia fuscolabialis 341

Abronia matudai 326

Abronia ochoterenai 326

Abronia oaxacae 337

Abronia taeniata graminea 349

Abronia taeniata taeniata 333

Aoris crepitans 359

Acrodytes inflata 331

Acrodytes modesta 325

Acrodytes spilomma 347

Adelophis copei 334

Adelphicos newmanorum 342

Adelphicos quadrivirgatus acutirostrum, 349
Adelphicos quadrivirgatus quadrivirgatus, 348

Adelphicos quadrivirgatus sargii 319

Adelphicos quadrivirgatus visoninus 320

Adelphicos veraepacis nigrilatus 326

Agalychnis callidrj-as 347

Agalychnis dacnicolor 328

Agalychnis moreletii 317

Agama torqiiata 329

Agkistrodon bilineatus bilineatus 317

Agkistrodon bilineatus taylori 346

Agkistrodon browni 332

Agkistrodon contortrix pictigaster 361

Agkistrodon mokeson pictigaster 361

Agkistrodon piscivorus leucostomus .... 360

Akleistops guatemalensis 319

Alligator chiapasius 326

Alligator lacordairei 316

Amastridium sapperi 317

Ambystoma amblycephalum 335

Ambystoma bombypellum 334

Ambystoma fluvinatum 327

Ambystoma granulosum 334

Ambystoma lacustris 335

Ambystoma ordinarium 335

Ambystoma rosaceum 327

Ambystoma schmidti 334

Ambystoma subsalsum 341

Ambystoma tigrinum proserpine 360

Ambystoma velasci 329

Ameiva f estiva edwardsii 317

Ameiva undulata amphigramma 350

Ameiva undulata dextra 332

Ameiva undulata gaigeae 353

Ameiva undulata hartwegi 326

Ameiva undulata parsa 319

Ameiva undulata podarga 346

Ameiva undulata sinistra 328

Ameiva undulata stuarti 326

Amei\a undulata thomasi 325

Ameiva undulata undulata 340

PAGE

Amj'da emoryi 361

Aneides lugubris lugubris 356

Anelytropsis papillosus 348

Anniella geronimensis 324

Anniella pulchra pulchra 357

Anniella texana 357

Anolis acutirostris 352

Anolis aureohis 351

Anolis baccatus 320

Anolis barkeri 346

Anolis beckeri 351

Anolis biporcatus 326

Anolis boidengerianus 337

Anolis capito 316

Anolis carneits 317

Anolis copei 317

Anolis cozumelae 342

Anolis cumingii 320

Anolis cymbops 340

Anolis damulus 315

Anolis dunni 331

Anolis gadovii 332

Anolis guntherii 320

Anolis heliactin 320

Anolis humilis uniformis 318

Anolis impetigosus 315

Anolis jacobi 351

Anolis kidderi 352

Anolis laeviventris 348

Anolis lemurinus bourgaei 348

Anolis limifrons rodriguezii 317

Anolis liogaster 332

Anolis megapholidotus 331

Anolis metallicus 320

Anolis milleri 338

Anolis nannodes 316

Anolis nebuloides 338

Anolis nebulosus 343

Anolis panamensis 320

Anolis pentaprion 364

Anolis petersii 317

Anolis petersii bivittatus 317

A7iolis rubigenosus 337

Anolis ruthveni 318

Anolis sagrei mayensis 325

Anolis sagrei sagrei 364

Anolis sallaei 348

Anolis schiedii 320

Anolis schmidti 328

Anolis sericeus 348

Anolis stuarti 317

Anolis sulcifrons 363

Anolis taylori 331

Anolis tropidonotus 348

(367)

368

The University Science Bulletin

PAGE

Anolis ustus 316

Anolis utowanae 343

Anolis wiegmanni 348

Anomalepis mexicana 320

Anota calidiarum 355

Anotheca coronata 316

Axolotes maculata 327

Arizona elegans eburnata 314, 356

Arizona elegans elegans 362

Arizona elegans exploita 314, 327

Arizona elegans pacata 314, 323

Arizona elegans philipi 314, 354

Arizona elegans occidentalis 356

Aristelliger georgeensis 316

Aristelliger irregularis 342

Barisia antauges 349

Barisia gadovii gadovii 332

Barisia gadovii levigata 338

Barisia imbricata ciliaris 328

Barisia imbricata imbricata 329

Barisia imbricata planifrons 337

Barisia levicollis 326

Barisia modesta 349

Barisia moreleti rafaeli 326

Barisia moreleti temporalis 325

Barisia rudicollis 334

Barisia viridiflava 338

Eascanion flagelhim frenatum 356

Bascanium semilineatus 353

Basilisciis nuchalis 320

Basiliscus vittatus 351

Bathysiredon dumerilii 335

Bathysiredon dumerillii queretarensis . . . 342

Batrachoseps attenuatus leucopus 323

Bipes biporus 322

Bipes canaliculatus 332

Bipes tridactylus 332

Boa divinoloquax mexicana 347

Bolitoglossa flaviventris 325

Bolitoglossa mexicana 339

Bolitoglossa moreleti 326

Bolitoglossa occidentalis 325

Bolitoglossa platydactyla 348

Bolitoglossa rufescens 349

Bolitoglossa yucatana 351

Bothriechis nummifera notata 317

Bothrops affinis 319

Bothrops atrox asper 320

Botlirops atrox dirus 363

Bothrops barbouri 332

Bothrops bicolor 319

Bothrops dunni 340

Bothrops godmani 319

Bothrops melanurus 341

Bothrops mexicanus 317

Bothrops nasutus 317

Bothrops nigroviridis aurifer 317

Bothrops nummifer nummifer 345

Bothrops nummifer veraecrucia 349

Bothrops schlegelii 363

Bothrops undulatus 350

PAGE

Bothrops yucatanicus 352

Bufo alvarius 355

Bufo angustipes 325

Bufo anomalus 329

Bufo argillaceous 328

Bufo boreas halophilus 357

Bufo californicus 357

Bufo canaliferus 339

Bufo coccifer 316

Bufo cognatus 357

Bufo compactilis compactilis 330

Bufo compactilis speciosus 361

Bufo cristatus 348

Bufo debilis 345

Bufo dipternus 358

Bufo eiteli 350

Bufo frontosus 353

Bufo gemmifer 331

Bufo horribilis 351

Bufo insidior 327

Eufo intermedius 364

Bufo kelloggi 343

Bufo lateralis 340

Bufo levifrons 330

Bufo niarmoreus 351

Bufo mazatlanensis 343

Bufo mexicanus 330

Bufo monksiae 330

Bufo nayaritensis 336

Bufo nebulifer 360

Bufo occidentalis 330

Bufo occipitalis 348

Bufo perplexus 332

Bufo punctatus 363

Bufo simus 320

Bufo sternosignatus 364

Bufo trachypus 351

Bufo valliceps 351

Caiman crocodilus fuscus 363

Callisaurus draconoides bogerti 343

Callisaurus draconoides brevipes 344

Callisaurus draconoides carmenensis .... 323

Callisaurus draconoides crinitus 321

Callisaurus draconoides draconoides 322

Callisaurus draconoides gabbii 321

Callisaurus draconoides inusitatus 345

Callisaurus draconoides ventralis 354

Callisaurus plasticus 321

Callisaurus rhodostictus 323

Callisaurus draconoides splendidus 323

Calopisma ciuinquevittatuni mexicana. . . . 350

Caretta bissa 315

Caretta caretta caretta 315

Caretta caretta gigas 314

Caretta remivaga 341

Caudisona atrox sonoraensis 344

Celestus atitlanensis 319

Celestus enneagranmius 348

Celestus rozellae 326

Centrolenella fleischnianni 316

Centrolenella vindissinia 331

Mexican Reptiles and Amphibians

369

PAGE

Chalcides sulcata 332 Cnemidophorus

Cham^esaura propus 332 Cnemidophorus

CheUorhina villarsii 334 Cnemidophorus

Chelonia agassizii 319 Cnemidophorus

Chelonia diissumieri 314 Cnemidophorus

Chelonia lachrymata 315 Cnemidophorus

Chelonia maculosa 315 Cnemidophorus

Chelonia mydas 315 Cnemidophorus

Chelonia pseudo-caret ta 315 Cnemidophorus

Chelonia pseudo-mydas 315 Cnemidophorus

Chelonia virgata 315 Cnemidophorus

Chelydra rossignonii 317 Cnemidophorus

Chelydra serpentina 358 Cnemidophorus

Chersodromus liebmanni 347 Cnemidophorus

Chersodromus nigricans 347 Cnemidophorus

Chilonieniscus cinctus 344 Cnemidophorus

Chilomeniscus punctatissimiis 324 Cnemidophorus

Chilonieniscus stramineus esterensis 321 Cnemidophorus

Chilomeniscus stramineus fasciatus 321 Cnemidophorus

Chilomeniscus stramineus stramineus.... 322 Cnemidophorus

Chionactis diasii 341 Cnemidophorus

Chionactis occipitalis annulatus 355 Cnemidophorus

Chionactis occipitalis palarostris 344 Cnemidophorus

Chiropterotriton arborea 333 Cnemidophorus

Chiropterotriton chiroptera 349 Cnemidophorus

Chiropterotriton chondrostega 333 Cnemidophorus

Chiropterotriton dimidiata 333 Cnemidophorus

Chiropterotriton lavae 349 Cnemidophorus

Chiropterotriton mosaueri 333 Cnemidophorus

Chiropterotriton multidentata 342 Cnemidophorus

Chiropterotriton terrestris 333 Cnemidophorus

Chiropterotriton xolocalcae 325 Cnemidophorus

Chrysemys picta bellii 358 Cnemidophorus

Churchillia bellona 327 Cnemidophorus

Cinostemum berendtianum 345 Cnemidophorus

Cinostemum brevigulare 320 Cnemidophorus

Cinostemum cobanum 317 Cnemidophorus

Cinostemum effeldtii 347 Cnemidophorus

Cinostemum henrici 359 Cnemidophorus

Cinostemum punctatum 354 Cnemidophorus

Cinostemum. rostellum 330 Cnemidophorus

Cinostemum. triliratum 339 Cnemidophorus

Claudius agassizii 345 Cnemidophorus

Claudius angustatus 345 Cnemidophorus

Claudius megalocephalus 345 Cnemidophorus

Claudius pictus 317 Cnemidophorus

Claudius severus 339 Cnemidophorus

Clelia clelia clelia 364 Cnemidophorus

Clelia clelia immaculata 334 Cnemidophorus

Clemmys marmorata pallida 356 Cnemidophorus

Cnemidophorus alfaronis 316 Cnemidophorus

Cnemidophorus arizonae 353 Cnemidophorus

Cnemidophorus bacatus 345 Cnemidophorus

Cnemidophorus bartolomas 321 Cnemidophorus

Cnemidophorus burti 344 Cnemidophorus

Cnemidophorus catalinensis 324 Cnemodophorus

Cnemidophorus ceralbensis 323 Cnemidophorus

Cnemidophorus communis copei 328 Cnemidophorus

Cnemidophorus commtmis occidentalis . . . 336 Cnemidophorus

Cnemidophorus costatus 336 Cnemidophorus

Cnemidophorus decemlineatiis 320 Cnemidophorus

23—90

PAGE

deppii cozumelus 342

deppii deppii 340

deppii lineatissimus 328

deppii oligoporus 350

dickersonae 324

disparilis 345

estebanensis 344

gracilis 355

grahami 362

gularis gularis obsoletus, 327

gularis gularis varus.... 327

gularis meeki 337

gularis sericevs 361

gidari^ velox 359

guttatus 361

guttatus guttatus 351

guttatus immutabilis. . . . 340

griftatus striatus 340

hyperythrus beldingi.... 323

hyperythrus caeruleus. . . 323

hyperythrus danheimae. . 324

hj'pery^hrus hyperythrus, 322

hyper>'thrus pictus .... 324

hyperythrus schmidti... 324

labialis 323

lativittis 338

marmoratus 361

maximus 322

melanostethus 355

mexicanus 341

mexicanus balsas 336

microlepidotus 340

motaguae 319

perplexus inornatus 337

perplexus perplexus 363

punctilineatus 345

sackii angusticeps 351

sackii australis 338

sackii bocourti 338

sackii communis 328

sackii gularis 361

sackii mariarum 336

sackii octolineatus 337

sackii sackii 336

sackii scalaris 327

semijasciatus 327

septemvittata 362

sexlineatus 360

sexlineatus tigris 327

stejnegeri 321

tesselatus 357

tigris aethiops 344

tigris canus 324

tigris celeripes 324

tigris martyris 344

tigris niultiscutatus. . . . 323

tigris mundtis 355

tigris rubidus 324

tigris tigris 363

undulatus 355

unicolor 337

370

The University Science Bulletin

PAGE

Cnemidophorus vandenburghi 323

Cnemidophorus variolosus 328

Cochliophagus tornieri 352

Coleonyx brevis 360

Coleonyx elegans elegans 316

Coleonyx elegans nemoralis 328

Coleonyx fasciatus 330

Coleonyx variegatus abbotti 357

Coleonyx variegatus bogerti 354

Coleonyx variegatus peninsularis 321

Coleonyx variegatus slevini 324

Coleonyx variegatus sonoriensis 344

Coleonyx variegatus variegatus 355

Coluber artzonae 362

Coluber constrictor stejnegerianus 361

Coluber novae hispaniae 349

Coluber oaxaca 337

Coluber sibon 349

Coluber striolatus 343

Coluber taeniatus girardi 362

Comastes quincunciatus 364

Coniophanes bipunctatus biseriatus 326

Coniophanes iissidens convergens 347

Coniophanes fissidens dispersus 331

Coniophanes fissidens fissidens 350

Coniophanes fissidens proterops 348

Coniophanes fissidens punctigularis 319

Coniophones imperialis clavatus 350

Coniophanes imperialis copei 340

Coniophanes imperialis imperialis 345

Coniophanes lateritius 334

Coniophanes meridanus 352

Coniophanes piceivittis 338

Coniophanes quinquevittatus 318

Coniophanes schmidti 352

Conophis lineatus concolor 352

Conophis lineatus lineatus 351

Conophis pulcher plagosus 326

Conophis pulcher similis 326

Conophis suniichrastii sumichrastii 334

Conophis vittatus viduus 340

Conophis vittatus vittatus 331

Conopsis biserialis 334

Conopsis nasus 330

Conopsis nasus heliae 342

Constrictor constrictor imperator 347

Constrictor constrictor sigma 336

Coronella multifasciata 357

Corythophanes chamaeleopsis 348

Corythophanes cristatus 349

Corythophanes hemandezii 348

Corythophanes mexicanus 348

Corythophanes percarinatus 318

Ctenosaura acanthura 346

Ctenosaura brevirostris 328

Ctenosaura (Cachryx) annectens 325

Ctenosaura completa 316

Ctenosaura cycluroides 351

Ctenosaura hemilopha 322

Ctenosaura insulana 323

Ctenosaura interrupta 322

PAGE

Ctenosaura multispinis 338

Ctenosaura parkeri 333

Ctenosaura pectinata 328

Ctenosaura similis similis 320

Ctenosaurus conspicuosa 344

Ctenosaurus teres brachylopha 343

Cristasaura mitrella 319

Crotalus atrox 361

Crotalus atrox elegans 323

Crotalus basiliscus basiliscus 328

Crotalus basiliscus oaxacus 314, 338

Crotalus cerastes laterorepens 356

Crotalus durissus durissus 348

Crotalus durissus totonacus 346

Crotalus enyo 322

Crotalus exsul 323

Crotalus gloydi 339

Crotalus gloydi lautus 314, 351

Crotalus goldmani 322

Crotalus jimenezii 334

Crotalus lepidus klauberi 353

Crotalus lepidus lepidus 362

Crotalus lugubris multimaculata 330

Crotalus lucasensis 321

Crotalus mitchellii mitchellii 322

Crotalus mitchellii pyrrhus 354

Crotalus molossus molossus 359

Crotalus molossus nigrescens 353

Crotalus omiltemanus 332

Crotalus polystictus 330

Crotalus pricei miquihuana 337

Crotalus pricei pricei 353

Crotalus ruber 356

Crotalus semicomutus 327

Crotalus scutulatus salvini 346

Crotalus scutulatus scutulatus 353

Crotalus stejnegeri 343

Crotalus tigris 354

Crotalus (tigris) palmeri 327

Crotalus tortugensis 325

Crotalus transversus 336

Crotalus triseriatus anahuacus 329

Crotalus triseriatus miquihuana 337

Crotalus triseriatus pricei 353

Crotalus triseriatus triseriatus 342

Crotalus viridis oreganus 360

Crotalus viridis viridis 358

Crotalus willardi 353

Crotaphytus coUaris baileyi 353

Crotaphytus copeii 321

Crotaphytus dickersonae 345

Crotaphytus fasciatus 359

Crotaphytus fasciolatus 321

Crotaphytus gambelii 355

Crotaphytus insularis 323

Crotaphytus reticulatus 363

Crocodylus acutus acutus 364

Crocodilvs americanus 351

Crocodilus bitcutatus 346

Crocodilus mexicanus 346

Crocodylus moreletii 318

Mexican Reptiles and Amphibians

371

PAGE

Crocodiliis pacificus 319

Cyclura denticulata 351

Cyclura semicristata 351

Cyclura shavfii 346

Cyclwra teres 346

Cystignathus frarjilis 340

Cystignathus microtis 335

Cystignathus perlaevis 339

Dermatemys abnormis 316

Dermochelys coriacea 315

Dendrophidion vinitor 318

Dermatemys mawii 346

Dermophis mexicanus mexicanus 347

Diadophis amabilis anthonyi 324

Diadophis amabilis similis 357

Diadophis dugesii 329

Diadophis regalis arizovae 354

Diadophis regalis blanchardi 361

Diadophis regalis laetus 354

Diadophis regalis regalis 344

Diaglena reticulata 340

Diaglena spatulata 343

Diemictylus kallerti 346

Diemictylus meridionalis 345

Diploglossus chalybeus 349

Dipsadomorus fasciatus 352

Dipsas bertholdi 315

Dipsas brevifacies 352

Dipsas brevis 320

Dipsas dimidiatus 320

Dipsas elegans 340

Dipsas gaigeae 328

Dipsas maxillaris 345

Dipsas sanniolus 352

Dipsosaurus carmenensis 323

Dipsosaurus catalinensis 324

Dipsosaurus dorsalis dorsalis 355

Dipsosaurus dorsalis lucasensis 322

Dipsosaurus dorsalis sonoriensis 344

Dirosema collare 347

Draconura bivittata 320

Dryadophis melanolomus melanolomus. . 352

Dryadophis melanolomus sleveni 336

Dryadophis melanolomus stuarti 331

Dryadophis melanolomus tehuanae 340

Dryadophis melanolomus veraecrucis. . . . 351

Drymarchon corals cleofae 336

Drymarchon corals erebennus 362

Drymarchon carats melanocercus 352

Drymarchon corais melanurus 352

Drymarchon corais obsoleta 362

Drymarchon corais orizabensis 350

Drymarchon corais rubidus 343

Drymarchon corais imicolor 325

Drymobius chloroticus 317

Drymobius margaritiferus fistulosus. . . . 336
Drymobius margaritiferus margaritiferus, 347
Drymobius margaritiferus occidentalis. . 319

Elapochrous deppei 348

Elaps corallinus crebripunctatus 350

PAGE

Elaps epistema 328

Elaps fulvius hypostema 328

Elaphe bairdi 362

Elaphe chlorosoma 334

Elaphe flavirufa flavirufa 352

Haphe flavirufa matudai 326

Elaphe laeta laeta 355

Elaphe rosaliae 323

Elaphe sclerotica 362

Elaphe subocularis 362

Elaphe triaspis 316

Elaphis pleurostictus 334

Eleutherodactylus alfredi 346

Eleutherodactylus augusti 330

Eleutherodactylus avocalis 340

Eleutherodactylus batrachylus 345

Eleutherodactylus beatae 349

Eleutherodactylus bolivari 334

Eleutherodactylus cactorum 341

Eleutherodactylus calcitrans 332

Eleutherodactylus conspicuus 318

Eleutherodactylus decoratus 348

Eleutherodactylus dorsoconcolor 351

Eleutherodactylus dunni 348

Eleutherodactylus hidalgoensis 333

Eleutherodactylus laticeps 351

Eleutherodactylus latrans 360

Eleutherodactylus longipes 320

Eleutherodactylus mexicanus 339

Eleutherodactylus macdougalli 338

Eleutherodactylus matudai 325

Eleutherodactylus natator 347

Eleutherodactylus occidentalis 353

Eleutherodactylus rhodopis 349

Eleutherodactylus rugulosus 337

Eleutherodactylus saltator 332

Eleutherodactylus spatulatus 347

Eleutherodactylus tarahumaraensis 327

Eleutherodactylus venustus 348

Eleutherodactylus vocalis 335

Elgaria cedrosensis 323

Elgaria kingii kingii 327

Elgaria kingii nobilis 359

Elgaria multicarinata nana 323

Elgaria multicarinata webbii 357

Elgaria paucicarinatas 323

Emys callirostris 346

Emys berardii 351

Emys grayi 319

Emys venusta 319

Engystoma mexicanum 341

Engystomops pustulosus 364

Ensatina croceater 356

Enulius sumichrasti 337

Enulius unicolor 331

Enyaliosaurus clarki 335

Enyaliosaurus defensor 352

Enyaliosaurus erythromelas 325

Enyaliosaurus quinquecarinatus 340

Epirhina tessellata 330

Eretmochelys imbricata 315

372

The University Science Bulletin

PAGE

Eietmochelys squaniata 315

Erythrolamprus aesculapii 315

Erythrolamprus grammophrj's 335

Erythrolamprus guentheri 320

Eudryas boddaerti mexicanus 351

Eumeces altamirani 335

Eumeces brevilineatus 360

Eumeces brevirostris 338

Eumeces callicephalus 330

Eumeces colimensis 328

Eumeces copei 334

Eumeces dicei 345

Eumeces dugesii 330

Eumeces gilberti rubricaudata 356

Eumeces humilis 343

Eumeces indubitus 336

Eumeces lagunensis 321, 322

Eumeces lynxe furcirostris 348

Eumeces lynxe lynxe 333

Eumeces multivirgatus 357

Eumeces obsoletus 363

Eumeces ochoterenai 332

Eumeces parviauriculatus 343

Eumeces parviilus 336

Eumeces quadrilineatiLS 356

Eumeces rovirosae 325

Eumeces schmidti 319

Eumeces schwartzei 325

Eumeces skiltonianus 360

Eumeces skiltonianus amblygrammus . . . . 355

Eumeces sumichrasti 350

Eumeces tetragrammus 345

Eumeces tetragrammus funebrosus 345

Eutaenia flaviiabris 329

Eutaenia insigniarum 329

Eutaenia macrostemma 329

Eutaenia pulchrilatus 330

Eutaenia rutiloris 342

Exelencophis nelsoni 336

Ficimia olivacea 350

Ficimia omata 352

Ficimia publia 352

Ficimia publia taylori 314, 350

Ficimia ramirez 338

Ficimia ruspator 332

Ficimia streckeri 362

Ficimia variegata 338

Gaigeia dontomasi 338

Gaigeia gaigeae 333

Gaigeia radula 339

Gaigeia sylvatica 333

Galedon annularis 349

Gambelia wislizenii wislizenii 359

Geagras longicaudatus 340

Geagras redimitus 340

Geatractus tecpanecus 332

Geoemyda areolata 318

Geoemyda pulcherrima incisa 316

Geoemyda pulcherrima pulcherrima 343

Geoemyda rubida 338

PAGE

Geophis anocularis 341

Geophis bicolor 329

Geophis blanchardi 346

Geophis cancellatus 325

Geophis chalybeus 320

Geophis dubius 340

Geophis dugesii 335

Geophis fuscus 348

Geophis isthmicus 340

Geophis latifrontalis 342

Geophis longiceps 350

Geophis maculiferus 335

Geophis mutitorques 333

Geophis multitorques yucatanicus 352

Geophis nasalis 318

Geophis omiltemana 332

Geophis petersii 335

Geophis rostralis 321

Geophis sallaei 338

Geoophis semiannulatus 328

Geophis semidoliatus 347

Geophis sieboldi 321

Gerrhonotus adspersus 334

Gerrhonotus bocourti 338

Gerrhonotus deppii digueti 341

Gerrhonotus lemniscatus 347

Gerrhonotus lichenigerus 329

Gerrhonotus liocephalus austrinus 325

Gerrhonotus liocephalus infernalis 363

Gerrhonotus liocephalus liocephalus.... 340

Gerrhonotus liocephalus loweryi 342

Gerrhonotus midtifasciatus 327

Gerrhonotus obscurus 338

Gerrhonotus olivaceus 329

Gerrhonotus liocephalus ophiurus 349

Gerrhonotus scincicauda ignavus 324

Gerrhonotus tessellatus 340

Glaphyrophis lateralis 346

Gonatodes fuscus 320

Goniodactylus braconnieri 363

Gopherus agassizii 356

Gopherus berlandieri 361

Gyalopion canum 354

Gyalopion desertorum 344

Gyalopion quadrangularis 343

Gymnodactylus coleonyx 318

Gyvinodactylus scapularis 318

Gymnophthalnius sumichrasti 341

Gynmopis multiplicata oaxacae 338

Helicops septemvittata 321

Heloderma Hernandesii 331

Heloderma horridum 331

Heloderma suspectum 354

Hcmidactylus exul 353

riemidactylus frenatus 314

Hemidactylus mabouia 364

Hemidactylus navarri 336

Hemidactylus turcicus turcicus 314

Hcmigenius variabilis 330

Herpetodryas incertus 316

Mexican Reptiles and Amphibians

373

PAGE

Heterodon nasicus kennerlyi 361

Holbrookia afEnis 363

Holbrookia bunkeri 327

Holbrookia maculata approximang 345

Holbrookia maculata dickersonae 327

Holbrookia maculata elegans 343

Holbrookia maculata flavilenta 359

Holbrookia maculata lacerata 362

Holbrookia maculata pulchra 353

Holbrookia maculata thermophila 344

Holbrookia propinqua 360

Holbrookia propinqua piperata 350

Holbrookia texana 361

Homalocranium boulengeri 348

Hydrops lubricus 350

Hyla affinis 354

Hyla arborescandens 346

Hyla arboricola 332

Hyla arenicolor 354

Hyla beltrani 326

Hyla bistincta 346

Hyla bromeliana 333

Hyla cardenasi 341

Hyla copii 361

Hyla crassa 320

Hyla curta 322

Hyla dalquesti 314, 349

Hyla dendroscarta 347

Hyla ebraccata 320

Hyla erythromma 331

Hyla euphorbiacea 347

Hyla eximia 329

Hyla forbesi 346

Hyla godmani 349

Hyla gracilipes 349

Hyla hazelae 339

Hyla holochlora 317

Hyla lafrentzi 329

Hyla leonard-schultzei 332

Hyla lichenosa 363

Hyla loquax 318

Hyla melanomma 331

Hyla microtis 341

Hyla miotympanum 348

Hyla muncolor 349

Hyla nigropunctata 348

Hyla pachyderma 350

Hyla phaeota 364

Hyla picta 348

Hyla pinorum 331

Hyla plicata 320

Hyla proboscidea 314, 349

Hyla regilla 363

Hyla regilla laticeps 322

Hyla rickardsi 350

Hyla robertmertensi 326

Hyla robertsorum 333

Hyla robustofemora 339

Hyla rozellae 325

Hyla smaragdina 335

Hyla smithi 336

PAGE

Hyla staufferi 349

Hyla taeniopus 348

Hyla underwoodi 316

Hyla vanvlietii 361

Hyla wrightorum 353

Hylella azteca 336

Hylella platycephala 339

Hylella sumichrasti 339

Hyliola digueti 336

Hylodes berkenbuschii 341

Hylodes plicatus 348

Hylodes sallaei 347

Hypopachus alboventer 336

Hypopachus caprimimus 331

Hypopachus cuneus cuneus 361

Hypopachus cuneus nigroreticulatus 325

Hypopachus maculatus 326

Hypopachus ovis 336

Hypopachus oxyrrhinus 343

Hypsiglena affinis 343

Hypsiglena chlorophea 354

Hypsiglena dunklei 345

Hypsiglena ochrorhynca janii 330

Hypsiglena ochrorhyncha klauberi. . .314, 324

Hypsiglena ochrorhyncha ochrorhyncha, 322

Hypsiglena ochrorhyncha texana. .. .314, 345
Hypsiglena ochrorhyncha

tortugaensis 314, 325

Hypsiglena ochrorhyncha venusta 323

Hypsiglena slevini 321

Hypsiglena torquata 320

Hypsiglena unaocularis 314, 329

Iguana (Ctenosaura) armata 346

Iguana (Ctenosaura) bellii 346

Iguana (Ctenosaura) lanceolata 346

Iguana iguana rhinolopha 347

Imantodes cenchoa leucomelas 349

Imantodes gemmistratus 316

Imantodes gracillimus 331

Imantodes latistratus 334

Imantodes splendidus luciodorsus 325

Imantodes splendidus oliveri 339

Imantodes splendidus splendidus 352

Imantodes tenuissimus 352

Kinostemon abaxillare 326

Kinosternon acutum 347

Kinosternon creaseri 351

Kinostemon cruentatum consors 342

Kinostemon cruentatum cruentatum 339

Kinosternon flavescens flavescens 362

Kinosternon flavescens stejnegeri 344

Kinosternon herrerai 349

Kinostemon hirtipes 343

Kinosternon integrum 331

Kinostemon leucostomum 347

Kinostemon mexicanum 339

Kinosternon sonoriense 354

I.acerta lumbricoides 332

Lacerta mexicana 332

Laemanctus alticoronatus 352

374

The University Science Bulletin

PAGE

Laemanctus deborrei 345

Laemanctus longipes 348

Laemanctus serxatus 349

Lampropeltis alterna 362

Lampropeltis califomiae 357

Lampropeltis catalinensis 324

Lampropeltis doliata annulata 345

Lampropeltis doliata arcifera 341

Lampropeltis doliata blanchardi 352

Lampropeltits doliata oligozona 337

Lampropeltis doliata polyzona 348

Lampropeltis doliata schmidti 336

Lampropeltis getulus boylii 355

Lampropeltis getulus califomiae 357

Lampropeltis getulus conjuncta 322

Lampropeltis getulus nitida 322

Lampropeltis getulus splendida 354

Lampropeltis getulus yumaensis 354

Lampropeltis knoblochi 327

Lampropeltis leonis 337

Lampropeltis mexicana 342

Lampropeltis nitida 322

Lampropeltis pyromelana 354

Lampropeltis ruthveni 335

Lampropeltis thayeri 345

Lam-propeltia triangulum annulata 345

Lampropeltis triangulum arcifera 341

Lampropeltis triangulum blanchardi 352

Lampropeltis triangulum nelsoni 330

Lampropeltis triangulum oligozona 337

Lampropeltis triangulum polyzona 348

Lampropeltis triangulum schmidti 336

Lampropeltis zonata agalma 321

Lampropeltis zonata herrerae 324

Lampropeltis zonata zonata 357

Lamprosaurus guttulatus 359

Leiolopisma assatum assatum 316

Leiolopisma assatum taylori 329

Leiolopisma cherriei cherriei 316

Leiolopisma cherriei ixbaac 352

Leiolopisma cherriei stuarti 350

Leiolopisma forbesorum 333

Leiolopisma gemmingeri 349

Leiolopisma laterals 358

Leiolopisma silvicolum 347

Lepidochelys kempii 358

Lepidochelys olivacea 315

Lepidophyma flavomaculatum

flavomaculatum 318

Lepidophyma smithi occulor 342

Lepidophyma smithii smithii 319

Lepidophyma smithii tehuanae 340

Lepidodactylus labialis 350

Lepidodactylus melanonotus 320

Lepidodactylus occidentalis 336

Leptodeira annulata polysticta 316

Leptodeira annulata septentrionalis 361

Leptodeira annulata taylori 350

Leptodeira bressoni 335

Leptodeira ephippiata 344

Leptodeira frenata 348

PAGE

Leptodeira guilleni 331

Leptodeira maculata 348

Leptodeira mystacina 340

Leptodeira personata 343

Leptodeira punctata 343

Leptodeira smithi 335

Leptodeira splendida 341

Leptodeira yucatanensis malleisi 325

Leptodeira yucatanensis yucatanensis. . . . 352

Leptognathus albocinctvs 343

Leptognathus bernoullii 319

Leptognathus cuculliceps 317

Leptognathus dumerili 349

Leptognathus leucostomus 352

Leptognathus subannulatus 352

Leptognathus semicinctus 317

Leptognathus sexscutatus 319

Leptognathus torquatus 352

Leptophis occidentalis praestans 318

Leptotyphlops bressoni 335

Leptotyphlops dugesii 328

Leptotyphlops dulcis 361

Leptotyphlops humilis cahuilae 357

Leptotyphlops humilig humilis 357

Leptotyphlops humilis segregus 361

Leptotyphlops humilis slevini 321

Leptotyphlops humilis tenuiculus 342

Leptotyphlops maximus 331

Leptotyphlops myopicus dissectus 359

Leptotyphlops myopicus myopicus 346

Leptotyphlops phenops bakewelli 328

Leptotyphlops phenops phenops 340

LAchanura myriolepis 321

Lichanura roseofusca gracia 356

Lichanura roseofusca roseofusca 321

Lichanura trivirgata 322

Liophis tricinctus 348

Liophis varia 350

Limnochelone micrura 346

Loxocemug bicolor 316

Loxocemus sumichrasti 340

Lysoptychus (Sceloporus) lateralis 361

Mabuia alliacea 316

Mabuya mabouya mabouya 364

Magnadigita macrinii 338

Magnadigita nigroflavescens 325

Magnadigita sulcata 320

Malaclemys terrapin littoralis 360

Manolepis putnami 336

Masticophis anthonyi 329

Masticophis aurigulus 322

Masticophis barbouri 324

Masticophis bilineatus 344

Masticophis flagellum lineatulus 327

Masticophis flagellum lineatus 343

Masticophis flagellum piceus 353

Masticophis flagellum testaceus 358

Masticophis flagellum variolosus 336

Masticophis rnentovarius mentovarius. . . 340

Masticophis lateralis 355

Mexican Reptiles and Amphibians

375

PAGE

Masticophis taeniatus australis 330

Masticophis taeniatus omatus 362

Masticophis taeniatus ruthveni 361

Masticophis taeniatus schotti 362

Masticophis taeniatus taeniatus 359

Microbatrachylus albolabris 347

Microbatrachylus hobartsmithi 335

Microbatrachylus imitator 325

Microbatrachylus lineatissimus 339

Microbatrachylus minimus 331

Microbatrachylus montanus 325

Microbatrachylus oaxacae 339

Microbatrachylus pygmaeus 350

Microhyla elegans 347

Microhyla olivacea 358

Microhyla mazatlanensis 343

Microhyla usta gado\'ii 339

Microhyla usta usta 333

Microphis quinqueliniativs 319

Micruroides euryxanthus 344

Micrurus bernadi 333

Micntriis browni 331

Micrurus ephippifer 340

Micrurus affinis affinis 347

Micrurus afRnis alienus 316

Micrurus afRnis apiatus 317

Micrurus affinis mayensis 352

Micrurus affinis stantoni 316

Micrurus diastema diastema 328

Micrurus diastema distans 326

Micrurus diastema michoacanensis 335

Micrurus elegans elegans 348

Micrurus fitzingeri fitzingeri 330

Micrurus fitzingeri microgalbineus 345

Micrurus fulvius tenere 361

Micrurus laticollaris 341

Micrurus latifasciata 319

Micrurus nigrocinctus browni 331

Micrurus nigrocinctus ovandoensis 326

Micrurus nigrocinctus wagneri 314, 325

Micrurus nigrocinctus zunilensis 319

Micrurus nuchalis nuchalis 339

Micrurus nuchalis taylori 331

Natrix erythrogaster transversa 300

Natrix rhombifera blanchardi 346

Natrix rhombifera rhombifera 360

Natrix valida celaeno 322

Natrix valida valida 330

Nerodia couchii 337

Ninia diademata diademata 350

Ninia diademata plorator 333

Ninia sebae morleyi 352

Ninia sebae sebae 351

Korops yucatanicus 352

Oedipina lineola 349

Opheodrys aestivus 360

Opheodrys mayae 352

Opisthiodon torquatus 348

Oreophis boulengeri 330

Oxybelis acuminatus 363

PAGE

Oxybelis aeneus auratus 340

Oxybelis fulgidus 352

Oxybelis microphthalmus 354

Oxybelis potosiensis 342

Oxyrhina defilippii 329

Oxyrhina (Exorhina) maculata 330

Oxyrhopus baileyi 350

Par\imolge townsendi 348

Pelamis platurus 315

Peropodum guatemalensis 318

Peropus mutilatus 315

Petalognathus multifasciatus 317

Petrosaurus repens 321

Petrosaurus thalassinus 322

Phrynonax guentheri 34C

Phrynosoma asio 328

Phrynosoma boucardi 333

Phrynosoma braconnieri 337

Phrynosoma bufonium 344

Phrjnosoma cerroense 323

Phrynosoma cornutum 358

Phrynosoma coronatum blainvillii 357

Phrynosoma coronatum coronatum 322

Phrynosoma coronatum frontale 356

Phrynosoma coronatimi jamesi 321

Phrynosoma ditmarsi 343

Phrynosoma douglassii brachycercum . . . . 330

Phrynosoma douglassii hemandezi 359

Phrynosoma harlanii 358

Phrynosoma m'callii 355

Phrynosoma modestum 359

Phrynosoma nelsoni 323

Phrynosoma orbiculare cortezii 349

Phrj'nosoma orbiculare dugesii 328

Phrynosoma ochoterenai 323

Phrynosoma orbiculare orbiculare 329

Phrynosoma planiceps 361

Phrynosoma platyrhinos goodei 344

Phrjnosoma platyrhinos platyrhinos. . . . 363

Phrynosoma regale 344

Phrynosoma schmidti 323

Phrynosoma spinimentum 340

Phrynosoma solare 355

Phrjmosoma taurus 341

Phrynosoma wiegmanni 329

Phyllodactylus bordai 332

Phyllodactylus delcampi 332

Phyllodactylus homolepidurus 344

Phyllodactylus lanei 332

Phyllodactylus magnatuberculatus 331

Phyllodactylus magnus 332

Phyllodactylus mentalis 315

Phyllodactylus muralis 340

Phyllodactylus pidcher 315

Phyllodactylus tuberculosus 321

Phyllodactylus unctus 322

Phyllodactylus xanti 322

Phyllorhynchus browni browni 354

Phyllorhynchus browni fortitus.... 314, 344
Phyllorhynchus decurtatus decurtatus. . . 322

376

The University Science Bulletin

PAGE

Phyllorhynehus decurtetus nubilis 354

Phyllorhynchus decurtatus perkinsi 356

Pituophis deppei hrevilineata 314, 335

Pituophis deppei deppei 334

Pituophis deppei jani 327

Pituophis deppei lineaticollis 348

Pituophis devppi pholidostictus 334

Pituophis catenifer affinis 359

Pituophis catenifer annectens 357

Pituophis catenifer bimaris 314, 323

Pituophis catenifer coronalis 314, 324

Pituophis catenifer deserticola 363

Pituophis catenifer fuliginosus 314

Pituophis catenifer fuliginatus 324

Pituophis catenifer insiilanus 314, 323

Pituophis catenifer rutilis 354

Pituophis catenifer sayi 358

Pituophis catenifer vertebralis 322

Pituophis haematois 322

Pituophis mexicanus 337

PlastosertjT bronni 316

Plectrohyla brachycephala 314, 339

Plectrohyla matudai 326

Plectrohyla sagorum 326

Plestiodon bellii ^^^

Pliocercus aequalis 317

Pliocercus andrewsi 352

Pliocercus bicolor 351

Pliocercus elapoides celatus 346

Pliocercus elapoides diastemus 317

Pliocercus elapoides elapoides 348

Pliocercus elapoides laticoUaris 345

Pliocercus elapoides schmidti 352

Pliocercus elapoides semicinctus 316

Pliocercus laticollaris 345

Procinura aemula 326

Psammophis flagivularis 360

Pseudemys floridana texana 361

Pseudemys scripta cataspila 346

Pseudemys scripta elegans 358

Pseudemys scripta gaigeae 361

Pseudemys scripta hiltoni 344

Pseudemys scripta omata 343

Pseudemys scripta nebulosa 321

Pseudemys umbra 319

Pseudodipsas fallax 364

Pseudoeurycea altamontana 336

Pseudoeurycea bellii 341

Pseudoeurycea cephalica cephalica 341

Pseudoeurycea cephalica manni 333

Pseudoeurycea cephalica rubrimembris. . 333

Pseudoeurycea cochranae 339

Pseudoeurycea gadovii 351

Pseudoeurycea galeanae 337

Pseudoeurycea gigantea 348

Pseudoeurycea leprosa 349

Pseudoeurycea melanomolga 351

Pseudoeurycea nigromaculata 347

Pseudoeurycea robertsi 334

Pseudoeurycea smithi 339

FAQE

Pseudoeurycea unguidentis 339

Pseudoficimia frontalis 328

Pseudoficimia hiltoni 314, 344

Pseudoficimia pulcherrima 331

Pseudoficimia pulchra 328

Pseudoleptodeira discolor 337

Pseudoleptodeira latifasciata 331

Pseustes poecilonotus argus 347

Pseustes poecilonotus poecilonotus 319

Pternohyla fodiens 343

Ptychohyla adipoventris 331

Ptychohyla bogerti 314, 339

Rana adtrita 329

Rana aurora draytoni 357

Rana catesbeiana 360

Rana forreri 343

Rana megapoda 333

Rana nioorei 314, 342

Rana montezumae 329

Rana montezumae concolor 330

Rana omiltemana 332

Rana palmipes 363

Rana pipiens 359

Rana pipiens austricola 314, 351

Rana pipiens trilobata 333

Rana pustulosa 330

Rana sierramadrensis 331

Rana tarahumarae 327

Rana trilobata 333

Rhabdosoma guttulatum 346

Rhadinaea aemula 332

Rhadinaea crassa 333

Rhadinaea decorata 350

Rhadinaea forbesi 351

Rhadinaea fulvivittis 350

Rhadinaea gaigeae 3*2

Rhadinaea hesperia baileyi 331

Rhadinaea hesperia hesperia 332

Rhadinaea hesperia hesperioides 334

Rhadinaea lachrymans 326

Rhadinaea laureata 329

Rhadinaea macdougalli 337

Rhadinaea marcellae 314, 343

Rhadinaea montana 337

Rhadinaea omiltemana 332

Rhadinaea quinquelineata 342

Rhadinaea taeniata 321

Rhadinaea vittata 350

Rhadinella schistosa 347

Rhinoclemmys mexicana 339

Rhinocheilus antonii clarus 356

Rhinocheilus lecontei antonii 336

Rhinocheilus lecontei lecontei 357

Rhinocheilus lecontei tessellatus 327

Rhinophrynus dorsalis 351

Rhinophrynus rostratus 340

Rhyacosiredon altamirani 329

Rhyacosiredon leorae 334

Rhyacosiredon rivularis 334

Rhyacosiredon zempoalaensis 336

Mexican Reptiles and Amphibians

377

PAGE

Salamandra togata 339 Sceloporus

Salvadora bairdii 330 Sceloporus

Salvadora bogerti 337 Sceloporus

Salvadora grahamiae 354 Sceloporus

Salvadora hexalepis celeris 343 Sceloporus

Salvadora hexalepis deserticolor 361 Sceloporus

Salvadora hexalepis hexalepis 354 Sceloporus

Salvadora hexalepis klauberi 314, 322 Sceloporus

Salvadora hexalepis virgultea 356 Sceloporus

Salvadora intermedia intermedia 331 Sceloporus

Salvadora intermedia richardi 341 Sceloporus

Salvadora lemniscata 331 Sceloporus

Salvadora lineata 362 Sceloporus

Salvadora mexicana 333 Sceloporus

Sator angustus 324 Sceloporus

Sator grandaevus 323 Sceloporus

Sauromalus ater 324 Sceloporus

Sauromalus australis 322 Sceloporus

Sauromalus hispidus 323 Sceloporus

Sauromalus klauberi 324 Sceloporus

Sauromalus obesus obesus 355 Sceloporus

Sauromalus obesus townsendi 345 Sceloporus

Sauromalus obesus tumidus 353 Sceloporus

Sauromalus slevini 324 Sceloporus

Sauromalus varius 344 Sceloporus

Scaphiodontophis albonuchalis 325 Sceloporus

Scaphiodontophis carpicinctus 318 Sceloporus

Scaphiodontophis cyclurus 347 Sceloporus

Scaphiodontophis nothus 350 Sceloporus

Scaphiodontophis sumichrasti 338 Sceloporus

Scaphiopus couchii 345 Sceloporus

Scaphiopus dugesii 329 Sceloporus

Scaphiopus hammondii 357 Sceloporus

Scaphiopus multiplicatus 329 Sceloporus

Scaphiopus rectifrenis 328 Sceloporus

Scaphiopiis varians 322 Sceloporus

Sceloporus aeneus aeneus 336 Sceloporus

Sceloporus aeneus bicanthalis 347 Sceloporus

Sceloponis asper 334 Sceloporus

Sceloporus bulleri 334 Sceloporus

Sceloporus carinatus 326 Sceloporus

Sceloporus cautus 827 Sceloporus

Sceloporus chrysostictus 352 Sceloporus

Sceloporus clarki boulengeri 343 Sceloporus

Sceloporus clarki clarki 354 Sceloporus

Sceloporus cochranae 341 Sceloporus

Sceloporus couchii 337 Sceloporus

Sceloporus cozumelae 342 Sceloporus

Sceloporus cyanogenus 337 Sceloporus

Sceloporus digueti 323 Sceloporus

Sceloporus dispar 347 Sceloporus

Sceloporus dugesii dugesii 328 Sceloporus

Sceloporus dugesii intermedius 335 Sceloporus

Sceloporus edwardtaylori 338 Sceloporus

Sceloporus ferrariperezi 330 Sceloporus

Sceloporus formosus formosus 346 Sceloporus

Sceloporus formosus scitulus 332 Sceloporus

Sceloporus fulvus 316 Sceloporus

Sceloporus gadovii 332 Sceloporus

Sceloporus goldmani 342 Sceloporus

Sceloporus graciosus vandenburgianus . . . 356 Sceloporus

PAGE

grammicus disparilis 362

grammicus grammicus 331

grammicus microlepidotus. . . . 329

guentheri 325

heterolepis 334

heterurus 349

horridus albiventris 336

horridus horridus 336

horridus oligoporus 328

humeralis 340

jalapae 348

jarrovii immucronatus 333

jarrovii jarrovii 353

jarrovii minor 353

jarrovii oberon 327

jarrovii sugillatus 335

lineolateralis 330

lundelli gaigeae 352

lundelli lundelli 316

maculosus 330

magister lineatulus 324

magister magister 355

magister monserratensis 324

magister rufidorsum 323

magister zosteromus 322

malachiticus acanthinus 319

malachiticus salvini 34h

malachiticus taeniocnemis . ... 317

megalepidurus 351

melanorhinus calligaster 331

melanorhinus melanorhinus. . . 340

melanorhinus stuarti 318

merriami annulatus 361

merriami merriami 363

mucronatus aureolus 346

mucronatus mucronatus 349

mucronatus omiltemanus 332

nelsoni 343

obscurus 336

occidentalis biseriatus 355

ochoterenae 332

olivaceus 362

orcutti licki 323

orcutti orcutti 356

ornatus caeruleus 328

ornatus ornatus 328

parvus parvus 337

parvus scutulatus 342

pictus 341

pilsbryi 342

pleurostictus 331

poinsettii 363

prezygus 325

pyrocephalus 328

ruhriventris 332

scalaris scalaris 329

scalaris slevini 353

scalaris unicanthalis 334

serrifer plioporus 347

serrifer serrifer 352

siniferus cupreus 341

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The University Science Bulletin

PAGE

Sceloporus siniferus siniferus 340

Sceloponis spinosus caeruleopunctatus . . . 339

Sceloporus spinosus spinosus 341

Sceloporus squamosus 318

Sceloporus stejnegeri 332

Sceloporus teapensis 345

Sceloporus torquatus binocularis 337

Sceloporus torquatus melanogaster 330

Sceloporus torquatus torquatus 329

Sceloporus undulatus consobrinus 360

Sceloporus undulatus virgatus 344

Sceloporus utiformis 328

Sceloporus variabilis marmoratus 361

Sceloporus variabilis smithi 340

Sceloporus variabilis variabilis 351

Sceloporus viviparus 349

Sceloporus westphalii 335

Schmidtophis rubriventris 314, 343

Scincus agilis 363

Scincus ventralis 329

Scolecophis scytalinus 345

Sibon nebulatus 349

Siredon harlanii 359

Siredon lermaensis 334

Siredon mexicanum 329

Siredon tigrina 329

Siren intermedia nettings 355

Sistrurus catenatus tergeminus 358

Sistrurus ravus 351

Smilisca baudinii baudinii 347

Smtlisca dauUnia 347

Sonora aequalis 321

Sonora bancroftae 322

Sonora episcopa 362

Soi}ora erythrura 332

Sonora michoacanensis michoacanensis . . 335

Sonora michoacanensis mutabilis 334

Sonora mosaueri 321

Sonora semiannulata blanchardi 361

Sonora semiannulata isozona 354

Sonora semiannulata linearis 355

Sonora semiannulata semiannulata 354

Sonora taylori 361

Spelerpes gibbicaudus 349

Spelerpes infuscatus 349

Spelerpes laticeps 347

Spelerpes orizabensis 349

Spelerpes punctatum 320

Sphaerodactylus glaucus glaucus 352

Sphaerodactylus lineolatus 320

Sphaerodactylus glaucus torquatus .... 338

Sphaerodactylus inomatus 340

Sphargis coriacea schlegelii 344

Sphenocalarmis lineolatus 343

Spilotes pullatus auribundu/S 349

Spilotes pullatus mexicanus 349

Spilotes salvini 319

Spilotes variabilis 349

Staurotypus biporcatus 318

Staurotypus marmoratus 339

Staurotypus salvinii 318

PAGE

Staurotypus triporcatus 346

Stenorrhina degenhardtii mexicana 347

Stenorrhina degenhardti quinquelineata . . 341

Stenorrhina freminvilli apiata 338

Stenorrhina freminvillii freminvillii 341

Stenorrhina freminvillii lactea 316

Storeria dekayi anomala 350

Storeria dekayi temporalineata 349

Storeria dekayi texana 362

Storeria hidalgoensis 333

Storeria storerioides 336

Streptophorus bifasciatus 350

Streptophorus sebae collaris 351

Streptosaurus mearnsi 356

Streptosaurus slevini 324

Swanka macidata 347

Symphimus leucostomus 338

Sympholis lippiens 334

Syrrhophus campi 361

Syrrhophus cystignathoides 350

Syrrhophus guttilatus 330

Syrrhophus latodactylug 337

Syrrhophus leprus 339

Syrrhophus modestus 328

Syrrhophus mystaceus 348

Syrrhophus nebulosus 326

Syrrhophus pipilans 332

Syrrhophus rubrimaculatus 325

Syrrhophus smithi 337

Syrrhophus verrucipes 333

Syrrhophus veiTuculatus 348

Tacliymenis m^lanocephala 341

Tantilla atriceps 337

Tantilla bimacuiata 343

Tantilla bocourti 330

Tantilla bocourti deviatrix 342

Tantilla bogerti 336

Tantilla calamarina 334

Tantilla canula 352

Tantilla coronadoi 331

Tantilla cuniculator 352

Tantilla deppei 321

Tantilla depressa 338

Tantilla deviatrix 342

Tantilla eiseni eiseni 355

Tantilla eiseni transmontanus 357

Tantilla hobartsmithi 344

Tantilla jani 317

Tantilla kimia 360

Tantilla martindelcampoi 331

Tantilla mexicana 321

Tantilla miniata 349

Tantilla moesta 318

Tantilla morgani 341

Tantilla nigriceps fumiceps 360

Tantilla nigriceps nigriceps 362

Tantilla phrenitica 347

Tantilla planiceps 322

Tantilla rubra 339

Tantilla shawi 314, 343

Mexican Reptiles and Amphibians

379

PAGE

Tantilla striata 338

Tantilla wilcoxi rubricata 337

Tantilla wilcoxi wilcoxi 353

Tantilla yaquia 327

Tantillita brevissima 326

Tantillita lintoni 318

Tapaya orbicularis longicaudata 329

Taricha klauberi 356

Terrapene coahuila 327

Terrapene goldmani 342

TeiTapene klauberi 344

Terrapene mexicana mexicana 346

Terrapene mexicana yucatana 351

Terrapene nelsoni 336

Terrapene omata 358

Testudo caouana 364

Testudo cephalo 360

Testudo viridis 360

Thalerophis diplotropis forreri 336

Thalerophis diplotropis diplotropis 340

Thalerophis mexicanus mexicanus 350

Thalerophis mexicanus yucatanensis 342

Thalorephis occidentalis praestans 318

Thamnophis angustirostris 328

Thamnophis arabdotus 342

Thamnophis chrysocephalus 350

Thamnophis cyrtopsis cyclides 330

Thamnophis digueti 332

Thamnophis eburatus 339

Thamnophis elegans hammondii 357

Thamnophis elegans hueyi 323

Thamnophis eques cyrtopsis 328

Thamnophis eques eques 351

Thamnophis eques postremus 335

Thamnophis errans 327

Thamnophis hammondii 357

Thamnophis macrostemma megalaps.... 354

Thamnophis marciana 355

Thamnophis melanogaster canescens. . . . 333
Thamnophis melanogaster linearis. . .314, 334
Thamnophis melanogaster melanogaster. . 330

Thamnophis ordinoides errans 327

Thamnophis ordinoides hueyi 323

Thamnophis ordinatus parietalis 359

Thamnophis phenax halophilus 333

Thamnophis phenax phenax 347

Thamnophis rozellae 326

Thomnophis rufipunctatus 353

Thamnophis ruthveni 340

Thamnophis sauritus chalceus 342

Thamnophis sauritus proximus 359

Thamnophis scalaris godmani 332

Thamnophis scalaris scalaris 348

Thamnophis scalaris scaliger 329

Thamnophis sirtalis parietalis 359

Thamnophis sirtalis proximus 359

Thamnophis stejnegeri 330

Thamnophis subcarinatus megalops 354

Thamnophis subcarinatus subcarinatus. . 334
Thamnophis sumichrasti cerebrosus .... 318
Thamnophis sumichrasti fulvu3 317

PAGE

Thamnophis sumichrasti praeocularis . ... 316
Thamnophis sumichrasti salvini.... 314, 317
Thamnophis sumichrasti sumichrasti.... 350

Thamnophis vicinus 335

Thecadactylus rapicaudus 352

Thorius dubitus 346

Thorius macdougalli 314, 338

Thorius minutissimus 314, 339

Thorius narisovalis 339

Thorius pennatulus 349

Thorius pulmonaris 339

Thorius troglodytes 346

Tomodactylus albolabris 331

Tomodactylus amulae 331

Tomodactylus angustidigitorum 335

Tomodactylus macrotympanum 333

Tomodactylus nitidus 341

Toluca amphisticha 314, 341

Toluca conica 339

Toluca lineata acuta 338

Toluca lineata lineata 329

Toluca lineata varians 346

Toluca lineata wetmorei 350

Toluca megalodon 339

Trimorphodon biscutatus biscutatus . . . . 340
Trimorphodon biscutatus semirutus .... 331

Trimorphodon collaris 351

Trimorphodon fasciolata 335

Trimorphodon forbesi 341

Trimorphodon lambda 344

Trimorphodon latifascia 341

Trimorphodon lyrophanes 322

Trimorphodon major 340

Trimorphodon paucimaculata 343

Trimorphodon tau 339

Trimorphodon upsilon 334

Trimorphodon vandenburghi 357

Trimorphodon vilkinsonii 326

Tropidodipsas annulifera 315

Tropidodipsas fasciata 352

Tropidodipsas fischeri 319

Tropidodipsas guerreroensis 331

Tropidodipsas macdougalli 340

Tropidodipsas occidentalis 328

Tropidodipsas philippii 343

Tropidodipsas sartorii armulatus 319

Tropidodipsas sartorii sartorii 349

Tropidonotus 7nesomelanus 330

Tropidonotus quadriserialis 343

Tropidonotus tephropleura 322

Tretanorhinus nigroluteus lateralis 316

Trimeresurus garciai 341

Triprion petasatus 351

Typhlops basimaculatus 350

Typhlops braminus 314

Typhlops longissimus 315

Typhlops microstomus 352

Typhlops perditus 350

Typhlops praelongus 347

Typhlops psittacus 321

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The University Science Bulletin

PAGE

Uma exsul 328

Uma notata cowlesi 344

Uma notata notata 355

Uma rufopunctata 355

Ungaliophis continentalis 318

Urosaurus auriculatus 329

Urosaurus bicarinatus anonymorphus . . . . 340

Urosaurus bicarinatus bicarinatus 336

Urosaurus bicarinatus nelsoni 338

Urosaurus bicarinatus tuberculatus 328

Urosaurus clarionensis 329

Urosaurus gadovii 333

Urosaurus graciosus 355

Urosaurus irregularis 320

Urosaurus microscutatus 322

Urosaurus nigricaudus 322

Urosaurus ornatus caeruleus 327

Urosaurus ornatus chiricahuae 353

Urosaurus ornatus linearis 344

Urosaurus ornatus ornatus 363

Urosaurus ornatus schmidti 362

Urosaurus ornatus schottii 344

Urosaurus ornatus symmetricus 355

Urosaurus unicus 326

Uta concinna 323

Uta gularis 344

Uta lateralis 343

PAGE

Uta mannophorus 323

Uta inartinensis 324

Uta nolascensis 345

Uta palmeri 344

Uta parva 321

Uta squamata 324

Uta stansburiana elegans 321

Uta stansburiana hesperis 356

Uta stansburiana stejnegeri 359

Uta stellata 324

Uta taylori 344

Verticaria esperitensis 324

Verticaria franciscensis 324

Virginia fasciata 319

Xantusia gilberti 322

Xantusia henshawi 357

Xantusia picta 356

Xantusia vigilis 356

Xenodon mexicanus 318

Xenosaurus fasciatus 348

Xenosaurus grandis 347

Xenosaurus newmanorum 342

Xenosaurus rackhami 317

Zamenis conirostris 345

Zamenis lateralis fidiginosus 324

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. II] March 20, 1950 [No. 9

Anomalies Found in a Series of Fetal and Newborn

Puppies

BY

Homer B. Latimer,
Department of Anatomy, University of Kansas

Abstract: Anomalies occurring in a series of 176 fetal and newborn puppies
are reported. The anomalies are grouped under three categories: hernias;
abnormally-shaped organs; and absence of organs. There are three cases of
hernia; five cases in which the thyroid and two in which the spleen has an
abnormal shape; there is one case in which the urinary bladder is absent, and
two in which one of the paired gonads is absent.

A series of 176 fetal and newborn puppies has been used in study-
ing the growth of the various organs, and several anomalies have
been observed and recorded. A record of these, and also the relative
incidence of the various types of anomalies, may be of interest,
hence this report is being presented.

There were 152 fetal and 24 newborn puppies in this series. They
were all obtained from the dogs used in the Physiology Department
and consequently were of all breeds, or rather, most of the dogs
were of no breed, being mongrels. None of the fetuses were taken
from dogs used in prolonged physiology experiments, which might
affect the development of the fetuses. The newborn puppies were
found at the animal house in the morning, having been born some
time during the night. These newborns were chloroformed and then
treated in the same manner as the fetuses. All of the specimens
were preserved in an adequate amount of 10 percent formalin after
the abdominal and the thoracic cavities had been filled with a 10
percent solution of formalin. Only enough formalin was injected
into these cavities, using a hypodermic syringe, to moderately fill
the cavity. Then an opening through the skin, fascia and brain

(381)

382 The University Science Bulletin

case was made at the top of the skull so that the fluid might enter
and preserve the brain and spinal cord. They were left in the
formalin solution for at least two months so that they might be
completely fixed and the evidence (Donaldson, '94; Hrdlicka, '06;
Kato, '38) seems to indicate that the changes, whatever they may
be, have become stabilized after this length of time in the pre-
servative.

There were 36 litters, with from 1 to 9 in each litter, or an average
of 4.89 puppies per litter. There were 6 litters of newborns and they
averaged just 4 puppies per litter. There were 86 of each sex and
4 were so small that their sex could not be determined positively in
the gross dissection. They ranged from 3 to 494 grams in body
weight and from 55 to 257 mm. in body length. The heaviest fetus
weighed 410 grams, but the longest specimen was a fetal dog.

Each specimen was removed from the formalin solution, all excess
liquid removed with paper toweling and then weighed on a labora-
tory balance to the nearest tenth of a gram for the larger specimens
and on a chemical balance to the nearest milligram for the smaller
specimens. These smaller specimens, as well as the organs, were
weighed in a glass stoppered weighing bottle to prevent drying dur-
ing the weighing. The lengths are the nose-anus lengths measured
along the back of the puppy. In the larger litters, where the fetuses
were crowded, they were much more curved than in those litters
with few and widely separated fetuses.

A regular routine of dissection was followed as described in an
earlier paper (Latimer and Corder, '48). When these anomalies
were found they were studied and recorded on the backs of the
record cards. It has seemed of some value to collect these and find
out some of the more common variations found in a series of fetal
and newborn dogs.

These anomalies may be grouped in three categories: (a) hernias;
(b) absence of organs and (c) abnormally shaped organs.

Hernias

In these 176 specimens three hernias were found, one umbilical,
one diaphragmatic and one through the muscular lacuna. This last,
a most unusual hernia has been described (Latimer, '48) and it will
not be described again. A diligent search of the literature has re-
vealed no case similar to this, described either in man or in any of
the domestic animals. This was a hernia of approximately one-
third of the length of the small intestine through the muscular la-
cuna. The herniated mass seemed to have followed the course of

Latimer: Anomalies of Newborn Puppies 383

the psoas major muscle out into the medial side of the thigh. An-
other hernia, also in a male fetus of 7.268 grams of body weight,
was a typical umbilical hernia. Only about two cm. of the small
intestine had herniated out through the umbilical opening whose
edges were rounded and not closely pressed on the hernia or the um-
bilical vessels. The third hernia was found in a 262.3 gram female
fetus. In this case the entire spleen had passed under the left lateral
lumbocostal arch into the left thoracic cavity. The vessels were
attached in a normal manner to the spleen and their other con-
nections were all normal except their length which was increased,
of course, due to the position of the spleen up in the thoracic cavity.
These last two types are found both in man and in the domestic
animals and they need no further discussion. Three cases of hernia
in 176 specimens gives an incidence of 1.7 percent for this series
of puppies.

One thing noticed in the handling and dissection of all of these
specimens was that the tension of the abdominal wall was much
greater than that of the thoracic wall. In both the fetuses and also
in the newborn pups the abdominal cavity was full while the thoracic
cavity was not filled completely. Even in the early newborn speci-
mens the lungs had not completely filled the thoracic cavities. In
these specimens the abdominal cavity appeared to be more than full
and in three of these there had been a herniation or protrusion of
a viscus from this cavity.

Abnormal Shape

The most common variation was the shape of the thyroid gland.
In these dogs there were usually two separate and distinct lobes
lying on either side of the trachea and in no way connected by
thyroid tissue. In five of these dogs, one male and four females, the
inferior or caudal ends of these paired lobes were connected by an
isthmus or a median mass of thyroid tissue thus resembling the
shape of the human thyroid gland. These specimens with the two
lobes of the thyroid connected by an isthmus ranged in body weight
from 8.624 to 220.5 grams of body weight. One female with a body
weight of 203.7 was a newborn puppy. Bradley ('43) describes the
thyroid as consisting of two lobes and "an isthmus of variable di-
mensions." He later says "Frequently the isthmus is absent." The
isthmus was present in but five of the 176 or in but 2.84 percent of
the specimens used in this study.

The rather frequent persistence of the old thyroglossal duct ex-
tending as far as the hyoid bone and found in man in 27 percent

384 The University Science Bulletin

of the cases (Gudernatsch, '42), was not seen in any of these dogs.
When one remembers the unpaired anlagen of the thyroid, it seems
that the human thyroid and these five anomalies are more primitive
than the type usually found in the dog. May we then call this an
atavistic anomaly?

Absence of an Organ

There are three cases of this type. In one male fetus weighing
13.6 grams there was no urinary bladder. Both ureters ended in a
mass of areolar tissue in the pelvis. The other two cases have one
of the paired gonads missing. A male fetus of 18.140 grams of body
weight had no left testis and a female of 20.6 grams of body weight
had no right ovary. In both of these cases, the organ of the other
side seemed to be perfectly normal in size, shape and location.

Anomalous Spleens

The last two anomalies are found in the spleen. The spleen in
the dog is an elongated and flattened organ, and very small in the
early fetus. One of these in a 97.5 gram male fetus had the dorsal
end bifurcated. The entire spleen was small, weighing but 0.1972
gram. The notch causing this bifurcated tip was not more than 3
mm. deep. In another male with a body weight of 56.0 grams, there
was a small accessory spleen. This spherical mass of splenic tissue
was about 2 mm. in diameter and located in the greater omentum.

All of these anomalies, with the exception of the first, have been
reported before but it seems worth-while to present these to give
some idea of the types of variations found in a series of fetal and
newborn dogs and the frequency of their occurrence. Had these
fetuses been permitted to go on to normal birth, most of these anom-
alies would not have prevented a normal adult life. The location
of the spleen in the thoracic cavity might not have made any dif-
ference in the normal life of this dog. The piece of the small in-
testine herniated out through the umbilicus might have been re-
tracted before birth, but the hernia through the muscular lacuna
would probably have been a serious thing for this newborn puppy.
Obviously the shape of the thyroid would have made no difference
in the postnatal life of these dogs, provided it secreted a normal
amount of thyroxine. The absence of one of the paired gonads
would have made no difference, for very likely the one persisting
gland would have hypertrophied and adequately cared for the need
of the animal. The absence of the urinary bladder would be a very
serious anomaly, had it not developed later before the birth of the
dog.

Latimer: Anomalies of Newborn Puppies 385

This absence of the urinary bladder and the hernia through the
muscular lacuna are probably the only cases which would have
seriously affected the postnatal life of these puppies. Thus, in only
two out of 176 fetuses and newborn puppies, or in only 1.1 percent
of this series, were there any anomalies which would in any way
have interfered with a normal postnatal life.

LITERATURE CITED

Bradley, O. C. 1943. Topographical anatomy of the dog. Macmillan Co.
New York.

Donaldson, H. H. 1894. Preliminary observations on some changes caused in
the nervous tissues by reagents commonly employed to harden them. J.
Morph., 9: 123-166.

GuDERNATSCH, J. F. 1942. Section on glands of internal secretion, Morris'
Human Anatomy. Blakiston Company, Philadelphia.

Hrdlicka, a. 1906. Brains and brain preservatives. Proc. U. S. Nat. Mus.
30: 245-320.

Kato H. 1938. Uber den Einfiuss der Fixerung auf des Hirngewicht. Folia
Anat. Japonica, 17: 237-295.

Latimer, H. B. 1948. A hernia through the muscular lacuna in a newborn
male puppy. Anat. Rec, 102: 439-444.

Latimer, H. B., and R. L. Corder. 1948. The growth of the digestive system
in the fetal dog. Growth, 12: 285-309.

24—90

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. II] March 20, 1950 [No. 10

The Bees of the Genus Proteriades
(Hymenoptera, Megachilidae)

BY
p. H. TiMBERLAKE and C. D. Michener,

Citrus Exp. Station, University of California, Riverside, California
and Department of Entomology, University of Kansas

Abstract: This paper is a revision of a geniis of bees, Proteriades, which is
known only from California and adjacent states. It is remarkable for having
peculiarly modified mouth parts which are used in obtaining pollen from the
flowers of Cryptantha. So far as known bees of this genus collect pollen from
no other plant. Proteriades is believed to have been derived from an An-
thocopa-\\ke ancestor.

The forms which have received the generic or subgeneric names Cephalapis
and Xerosmia are here included in Proteriades. A total of twenty-two species
are recognized, fourteen of which are described as new. In addition, three
new subspecific names are proposed.

Proteriades is a genus of small megachilid bees, allied to Hoplitis
and Anthocopa, and remarkable for having rather numerous species
in spite of its limited distribution and for having specialized pollen
collecting apparatus enabling its members to obtain pollen from
flowers of Cryptantha, upon which it seems to be entirely dependent.

Distribution
All the known species of the genus except P. incanescens (Cock-
erell) have been found only in California and occur from near sea
level to 9,500 feet altitude in the mountains. P. incanescens ranges
from eastern California into Nevada and Arizona. Presumably
other species will also be found to occur in these states. This dis-
tributional pattern parallels that of certain other megachilid bees
(see Michener, 1943, 1944a) which for some reason do not range
nearly as far eastward and southward as does the desert habitat in
which they occur in California.

(387)

388 The University Science Bulletin

Of the twenty-two species now recognized, fifteen are known in
both sexes, with the sexes in most cases certainly allocated, but in
a few cases a little uncertainly. Five species are known only in the
female sex, and two only in the male sex. Several species are
known to be widely distributed in California, while others remain
known from only one or a few localities. One of us (Timberlake)
has collected nine species, over one-third of the known species of
the genus, on or in the immediate vicinity of the grounds of the
Citrus Experiment Station in Riverside, California. This was done
over a period of more than twenty years, and one species {similis)
was found but once and another (seminigra) only during one year.
Thus any locality where suitable conditions exist may continue to
yield additional species for many years. We may have a good
proportion of the species, but there certainly must be others still
unseen. It is evident that much more material needs to be collected,
and that many years may elapse before a full understanding of the
genus is achieved.

Pollen Collecting

It seems unquestionably true that all species of Proteriades col-
lect pollen exclusively from flowers of the boraginaceous genus
Cryptantha. The mouth parts (galeae and labial palpi) are provided
with stiff curled hairs, especially in the female. These serve to
pull the pollen from the stamens which are hidden in the throats
of the flowers. These flowers are too small to permit the bees to
enter bodily and collect pollen in the usual way. From the curled
or hooked hairs of the mouth parts the pollen is transferred for
carrying to the hairs of the scopa on the under side of the abdomen.

Thus far there is no evidence whatever to suggest that the dif-
ferent species of Proteriades are restricted to different species of
Cryptantha. Several species of Proteriades have been collected on
more than one species of Cryptantha, and the indications are that
a species of Proteriades will utilize whatever Cryptantha species are
available in the neighborhood. All eight species of Proteriades
found at Riverside were on Cryptantha intermedia.

There is no evident relationship between speciation in Cryptantha
and that in Proteriades. Not only are the Proteriades species not
restricted to particular species of Cryptantha, but the distribution
of the plant is far wider than that of the bee, extending eastward to
the Great Plains and southward far into Mexico and even to South
America. In this connection it should be pointed out that in the
vicinity of Riverside, where eight species of Proteriades are com-

Bees of the Genus Proteriades 389

peting with one another, visiting the same flowers at the same season
in the same place, the season of flight of the bees does not even
coincide with the main bloom of the Cryptantha plants. These plants
appear in great numbers, especially in bare and disturbed soils, and
are at the height of their bloom in March. Many of them may dry
up before the end of that month. Other plants, more favorably
situated, as on a north slope or at the edge of a boulder or wherever
the soil retains its moisture longer than in exposed places, may
bloom through April and May into June. It is mainly such plants
which support the Proteriades populations.

However, at high altitudes in the Inyo Mountains the Proteriades
are on the wing a few days before the first Cryptantha flowers are
in bloom. Here the situation is wholly different from that in coastal
southern California (Riverside) for in these high desert ranges the
season is extremely short and probably no Cryptantha plant sur-
vives the aridity to bloom for more than a few weeks.

While collecting pollen at flowers the females are easy to collect,
in comparison with the males, and it is partly on account of this
that as many as five species are kno^\Ti only from the female sex,
The females also undoubtedly have a much longer season of flight.
The males, on the other hand, have the habit of sunning themselves
on the ground, or on a convenient stone or stick, from which they
make periodic sorties over the Cryptantha flowers apparently in
search of mates, only occasionally lingering at the flowers to feed.

The nesting habits of only P. xerophila (Cockerell) have been
observed, but presumably all the species nest in the ground. As
reported by Michener (1943), P. xerophila was reared from old
nests of Anthophora linsleyi Timberlake which were dug from the
ground by Doctor G. E. Bohart.

Phylogeny

From our meager knowledge it is impossible to draw a reliable
phylogenetic tree of Proteriades, but it is possible to suggest the
relationships of certain species. The genus apparently evolved
from an Anthocopa-like ancestor, as it agrees with that genus in the
robust form, in the clypeus usually produced well over the base
of the labrum, and in having the inner surface of the hind coxae
flattened and carinate along the inner ventral margin. The de-
parture from an Anthocopa-like structure is in having specialized,
stiff, hooked hairs in the mouth parts and in having the base of the
first metasomal tergum much less basinlike and more sulcate me-
dially as in the genus Hoplitis although to a less degree.

390 The University Science Bulletin

As the bulk of the species of Anthocopa and Hoplitis, as well as
of other related genera such as Osmia, have five-segmented maxil-
lary palpi, it is clear that this is a primitive characteristic. It is
retained in only two species of Proteriades, remotula and incanescens.
These species, although specialized in certain respects, appear to
constitute the most primitive group of Proteriades, which we will
call the remotula group. It is perhaps significant that the group is
relatively widespread, occurring from northernmost California to
the southern part of the state and eastward at least to Southern
Nevada and the Grand Canyon region of Arizona. The male of
remotula is remarkable in having very few hooked hairs on the
mouth parts, and these but little hooked. This may be another
primitive characteristic. Unfortunately the male of incanescens is
unknown.

No doubt derived from the remotula group is a large group, the
bulk of the genus, which may be called the tristis group. It differs
from the remotula group principally in having four or less seg-
ments in the maxillary palpi. The species included are basingeri,
bidenticauda, caudex, cryptanthae, hamulicornis, nanula, nigra,
palmarum; pygmaea, reducta, seminigra, similis, tristis, truicauda,
and xerophila. Specialization along several different lines is evi-
dent in this group. For example, reduction in the maxillary palpi
to two segments occurs in reducta, reduction in size in nanida and
pygmaea. The males of several species show special modifications;
for example, the acute antennae of hamulicornis, the ventral pro-
tuberance of the base of the abdomen of caudex and cryptanthae,
the long scoop-shaped apical tergum of truicauda, and the bidentate
apical tergum of bidenticauda. The last structure is similar to that
of deserticola, a member of the semirubra group, but there seems
to be no close relationship between bidenticauda and deserticola,
the bidentate tergum having arisen independently in the two groups.

P. xerophila differs from other members of the tristis group prin-
cipally by its large size and the tridentate seventh metasomal tergum
of the male. The sixth sternum has a pair of lobes, one on each
side of the short median fold. As these differences are no more
striking than the differences between certain members of the tristis
group, xerophila is included in that group. It is the type species
of Xerosmia, a name which might well be used in a subgeneric sense
for all members of the remotula and tristis groups.

The semirugra group is characterized by very distinctive females
but has males scarcely distinguishable from those of the tristis

Bees of the Genus Proteriades 391

group. It is no doubt derived from the tristis group. In the fe-
males the mandibles are thick, not gradually narrowed subbasally
as in the remotula and tristis groups. The clypeus of the female
is thickened, either short and but little overhanging the labrum or
produced into a pair of fingerlike projections. In the male the sec-
ond metasomal sternum has a broad transverse subapical ridge which
extends uniformly across the segment. In members of the tristis
and remotula groups this ridge, if present at all, is interrupted
medially. Species included in the semiruhra group are evansv, de-
serticola, and semiruhra. P. boharti is tentatively included al-
though unknown in the female. In the event of a subgeneric division
of Proteriades, this group would be called Proteriades s. str.

The last group contains the single species jacintana, which is re-
markable for its elongate form, broad genal areas, keeled clypeus
in both sexes, the long nearly straight margin between the second
and third mandibular teeth of the female, etc. This species may
be placed under the subgeneric name Cephalapis, but as it is merely
one more, highly specialized derivitive of the tristis group, the use
of the subgeneric name seems unnecessary, unless Xerosmia is also
utilized in a subgeneric sense for the tristis and remotula groups.

Systematic Treatment
The genus Proteriades was established by Titus (1904) for the
male of a California bee described by Cockerell (1898) as Heriades
semiruhra. Other species that belong here are Chelostoma {Cephala-
pis) jacintanum Cockerell (1910), Osmia remotula Cockerell (1910),
Hoplitina incanescens Cockerell (1922), Osmia xerophila and 0.
palmarum Cockerell (1935), and Proteriades evansi and P. tristis
Michener (1936). Cephalapis Cockerell, which was described as a
subgenus of Chelostoma, was based on a very distinctive species of
Proteriades and possibly deserves recognition as a subgenus in its
new position. In 1943 Michener erected the group Xerosmia for
Osmia xerophila Cockerell and placed it as a subgenus in the genus
Anthocopa. After reconsideration in the light of the numerous addi-
tional species of Proteriades now known, Xerosmia is removed to
Proteriades, with which it agrees in every way except for certain
specialized male characters. At that time it was believed (Michener
1941, 1943, 1944) that Proteriades was more closely related to
Hoplitis than to Anthocopa, and it was the AnthocopaA^ko. char-
acteristics of xerophila that lead to its inclusion in Anthocopa as a
subgenus. It is now apparent that other Proteriades share these
characteristics, and xerophila must be placed with Proteriades,

392 The University Science Bulletin

which, however, could logically be regarded as a subgenus of
Anthocopa.

The following is the synonymy of Proteriades:

Proteriades Titus

Proteriades Titus, 1904, Jour. New York Ent. Soc, 12: 25; Cockerell, 1906,
Bull. Amer. Mus. Nat. Hist., 22: 445; Michener, 1936, Amer. Mus. Novitates,
875: 28; Sandhouse, 1939, Mem. Ent. Soc. Washington, 1:2; Michener, 1941,
Amer. Midland Nat., 26: 160; Sandhouse. 1943, Proc. U. S. Nat. Mus., 92: 591;
Michener, 1944, Amer. Nat., 78: 262; Michener, 1944, Bull. Amer. Mus. Nat.
Hist., 82 : 263.

Type: Heriadcs semiruba Cockerell (monobasic).

Cephalapk Cockerell. 1910, Ann. Mag. Nat. Hist., (8)5: 23; Sandhouse, 1943,
Proc. U. S. Nat. Mus., 92: 535.

Type: Chelosloma {Cephalapis) jacintanum Cockerell (monobasic and orig-
inal designation),

Xerosmm Michener. 1943, Ann. Ent. Soc. Amer., 36: 81; Michener, 1944,
Bull. Amer. Mus. Nat. Hist., 82: 264.

Type: Osmia xerophila Cockerell, by original designation.

As elsewhere explained, the genus can be recognized by the
hooked or wavy hairs of the galeae and labial palpi. The first ab-
dominal segment is intermediate in structure between that of Antho-
copa and that of Hoplitis, being sulcate but more broadly so than
usual in Hoplitis. Almost all of the species are small, black, with
the abdomen partly red.

As indicated for each of the new species, holotypes are in the
collections of the Citrus Experiment Station of the University of
California, the California Academy of Sciences or in the Snow En-
tomological Collections of the University of Kansas. Paratypes of
each species, wherever posible, are divided between these collections
and where series of paratypes exist some have been sent to the
American Museum of Natural History and the United States Na-
tional Museum.

Key to the Species of Proteriades

FEMALES

1. Clypeus with longitudinal median carinate ridge; mandible with apical tooth basad

of which is a notch, followed by a long straight margin (fig. 7) ; genal area

twice as broad as eye seen from side jacintana

Clypeus without longitudinal ridge and without carina except in P. palmarum;

mandible tridentate (fig. 9) ; genal areas at most but little broader than eye. . . 2

2. Clypeus produced on each side (next to base of mandibles) as much as, or more

than, in the middle, and not overhanging base of labrum or doing so only
medially 3

Clypeus distinctly more produced in middle than at the sides and overhanging base

of labrum 4

8. Side of clypeal margin more produced than the middle, which is much elevated
and perpendicularly declivous, the margin strongly arched as seen from beneath
and leaving base of labrum fully exposed ; mandibular teeth nearly equal . . . evansi

Clypeal margin about equally produced at sides and in the middle, with a short
median process that is moderately elevated over base of labrum ; mandibles
with inner apical tooth reduced to an angle and much shorter than the two
acute outer teeth deserticola

Bees of the Genus Proteriades 393

4. Clypeus truncate or rounded at apex; mandibles not broadened medially 5

Median lobe of clypeus strongly produced and with two fingerlike divergent proc-
esses between which is a small acute tooth ; mandibles very broad at middle,
deeply excavated on inner margin toward the base, and a little narrowed to
apex semirubra

5. Clypeal margin in middle more or less thin-edged and strongly projecting over

base of labruni, not or feebly crenulate 6

Clypeal truncation thick-edged, crenulate with four or six small blunt teeth, and

but little produced over base of labrum cryptanthae

6. Upper half or two-thirds of clypeus strongly swollen and shining and nearly im-

punctate medially ; maxillary palpi five -segmented 7

Clypeus more uniformly convex with the punctures moderately dense to crowded
, (a median impunctate line in 'palmarum) ; maxillary palpi with two to four

segments 8

7. Clypeal margin broadly rounded with no distinct angles demarking truncation ;

mesoscutum with punctures not coarser than those of vertex remotula

Clypeal margin with a clearly defined truncation ; mesoscutum with punctures

coarser than those of vertex incanescens

8. Abdomen black, or at most with a small amount of red on first tergum and sides

of second ; terga always with distinct white hair-bands ; clypeal margin broadly

truncate, or emarginate-truncate 9

Abdomen with mucli red on the three basal segments, the hair-bands well de-
veloped or not ; clypeal margin usually less distinctly truncate, sometimes
plainly rounded 12

9. Without red on abdomen (desert species) 10

Sides of first one or two segments of abdomen red (coastal and montane species),

tristis

10. Smaller species, about 6 mm. long ; distance between posterior ocelli subequal to

distance from one of them to eye margin 11

Robust species, usually about 8 mm. long, rarely only 6.5 mm.; distance between

posterior ocelli less than distance from one of them to eye margin xerophila

11. Clypeus strongly convex above the middle with longitudinal median raised im-

punctate line; emargination between second and third mandibular teeth ar-
cuate palmarum

Clypeus less strongly convex, without such a line or with it weakly developed ;

emargination between second and third mandibular teeth angular nigra

12. Clypeal margin broadly rounded, with a very obtuse sublateral angulation far to

each side ; small species about 3.0 to 4.6 mm. long 13

Clypeal margin more or less distinctly truncate in middle, that is with rounded
angles demarking truncation in addition to sublateral angles ; length about
5 mm 15

13. Clypeus very densely punctured; prominences in front of ocelli shining and dis-

tinctly more sparsely punctate than vertex and rest of supraantennal area. . sim,ilis
Clypeus less densely punctured, with interspaces between punctures; prominences
in front of ocelli not or scarcely more sparsely punctate than vertex and rest
of supraantennal area 14

14. Small desert species, with pubescence white and unusually abundant on face,

cheeks, sides of thorax and margins of mesoscutum ; hair bands of abdomen
broad and dense, the disk of last three terga covered with white hair ; punc-
tures of mesoscutum about a puncture-width apart ; length usually not over

4 mm pygmaea

Slightly larger montane and coastal species with pubescence less clear white, less
dense on face and thorax; disk of fourth tergum well exposed, that of fifth
and sixth segments with sparse white hair; punctures of mesoscutum mostly
less than a puncture width apart ; length 3.5 to 4.5 mm nanula

15. Apical margin of clypeus not distinctly concavely arcuate on each side of the

truncation 16

Apical margin of clypeus rather distinctly but shallowly emarginate on each side
of the rather narrow truncation, the sublateral angulation strong but obtuse.
(Punctures of clypeus close, those of mesoscutum rather coarse and almost uni-
formly less than a puncture width apart) caudex

394 The University Science Bulletin

16. Superior part of supraantennal area (frons), especially the slightly prominent areas

just in front of the lateral ocelli, having the punctures more or less separated. . 17
Supraantennal area uniformly and very closely punctured, the surface appearing

more dullish 19

17. First three metasomal terga and generally most of the fourth, red; abdominal

bands weak; somewhat smaller species, about 4.5 to 5.5 mm. long 18

Abdomen black, the sides of all terga red, but decreasingly so toward apex ; apical
margins of first five terga conspicuously pallid, and the first four each with a
dense white hair-band. Length 5.5 to 6 mm. (Sublateral angulations of cly-
peal margin very obtuse, so that the margin appears rather broadly rounded.),

truicauda
IS. Maxillary palpi short, indistinctly two- or three-segmented ; apical truncation of
clypeus rather distinct and about equal to the oblique sides, which have the

sublateral angulation very obtuse reducta

Maxillary palpi four-segmented and usually distinctly longer than in reducta;
clypeal margin usually distinctly although minutely notched (the notch bear-
ing a seta) just inside the lateral angulations, so that the latter are produced

to form small blunt teeth bidenticauda

19. Clypeal truncation distinctly less in width than the oblique sides of the apical
margin; punctures of clypeus at least a puncture-width apart; punctures of
mesoscutum close, but rather more separated than those of frons; first three

terga, and sometimes a large part of fourth, red hamulicornis

Clypeal truncation about equaling width of oblique sides of the apical margin;
punctures of clypeus rather less than a puncture-width apart; punctures of
mesoscutum about equaling those of frons in density ; basal terga more or less
black in middle, the first sometimes only narrowly so, the fourth either en-
tirely black or red at sides seminigra

MALES

1. Mandible longer than front tibia, with subapical dorsal angle so that it is vir-

tually tridentate; clypeus with longitudinal median carina; emargination be-
tween lobes of ninth tergum deeply U-shaped jacintana

Mandible shorter than front tibia, bidentate ; clypeus without longitudinal carina ;
ninth tergum, if bilobed, with emargination between lobes broad and shallow
(except in remoiula) 2

2. Seventh metasomal tergum tridentate; sixth sternum with two small but strongly

projecting lobes on either side of base of median fold xerophila

Seventh tergum truncate, bidentate or bilobed ; fifth sternum without such

lobes 3

S. Seventh metasomal tergum bidentate at apex, the teeth small, separated by at

least four times the width of a tooth 4

Seventh tergum truncate at apex, or notched medially, the teeth or lobes in that

case at least half as broad as the space between them 5

4. Teeth of seventh tergum separated by a creseentiform emargination; second ster-

num with a broad, subapical, transverse ridge ; fifth sternum fringed ; abdomen

with broad white apical hair-bands deserticola

Teeth of seventh tergum separated by a quadrate emargination ; second sternum
with an oblique swelling on each side subapically; fifth sternum not fringed,
its posterior margin broadly transparent, without hairs ; abdomen without
hair-bands bidenticauda

5. Flagellum rounded at apex; posterior coxae not concave mesially, not spined 6

Apical segment of flagellum produced beneath at apex into a fine, sharp hook ;

posterior coxae concave mesially, with long sharp spine arising from inner side

at apex hamulicornis

G. First sternum with a large hornlike swelling at apex 7

First sternum with at most a low median protuberance at apex 8

7. Process of first sternum narrowing to transverse truncate, thin-edged apex, its pos-
terior face flat and perpendicular; seventh tergum truncate at apex. . . cryptanthae
Process of first sternum thick, trunklike, ending bluntly, its posterior face convex ;
second to fifth sterna each with an apical fringe; seventh tergum strongly
emarginate at apex, the two lobes a little less wide than the median emargi-
nation caudex

Bees of the Genus Proteriades 395

8. Seventh tergum no longer than preceding one; second sternum without a pro-

tuberance on cacli side 9

Seventh tergum much lengthened, the apical portion convex above, concave be-
neath, broadly truncate at apex, its lateral margins nearly straight and un-
armed; second sternum with a protuberance on each side subapically. . . . truicauda

9. Impunctate margin of clypeus broadly emarginate, with small median protuber-

ance or toolh ; angles demarking clypeal truncation acute semirubra

Margin of clypeus emarginate or not but without median tooth, often crenulate;

angles demarking ; lypeal truncation obtuse 10

10. Posterior coxae each with irmer carina elevated to form a lamella which is ab-

ruptly terminated at its apex; flagellar segments longer than broad boharti

Posterior coxal carinae normal ; flagellar segments shorter, at least subapical ones

broader than long or rarely as long as broad 11

11. First sternum nearly nude, or thinly and uniformly hairy 12

First sternum with a triangular medio-apical area densely covered with white hair,

which also fringes the margin. (Length at least 5 mm. ; middle coxae not
toothed.) reducta

12. Maxillary palpi five-segmented; seventh tergum with a deep almost semicircular

emargination separating lobes which are about half as wide as emargination,

re7notuIa

Maxillary palpi three- or four-segmented; seventh tergum truncate or with shallow

emargination separating lobes which are at least as wide as emargination 13

13. Larger species, 5 to 7 mm. long; middle coxae each with a ventral angle or tooth

in front of base of trochanter ; ^ lateral margins of seventh tergum strongly

angulated ; teeth of mandibles equal or subequal (except in seminigra) 14

Small species, about 3 to 4.5 mm. long; middle coxae not toothed; lateral mar-
gins of seventh tergum very obtusely and weakly angulated ; outer tooth of
mandibles much longer than inner tooth 16

14. First sternum with a definite shining medio-apical protuberance ; abdomen

almost wholly red, infuscated apically basingeri

First sternum nearly evenly and weakly convex across the apex ; abdomen with

black dorsally on most or all of the terga 15

15. Seventh tergum with hardly a trace of a median notch ; first three or more meta-

somal terga with red seminigra

Seventh tergum definitely notched to form two rounded lobes; red confined to

sides of first two metasomal terga tristis

16. Carina of inner ventral angle of posterior coxa weak, absent distally where coxa is

beveled ; distance from posterior ocellus to extreme posterior margin of vertex
little more than diameter of ocellus ; genal areas little more than half as wide
as eyes seen from side; pubescence dense and brilliantly white, obscuring sculp-
turing of face up nearly to level of ocelli .' pygmaea

Carina of inner ventral angle of posterior coxa conspicuous and complete, coxa not
beveled ; distance from posterior ocellus to extreme posterior margin of vertex
nearly twice diameter of ocellus; genal areas much more than half as wide as
eyes seen from side; pubescence dull white, not obscuring facial sculpturing
above level of antennal bases nanula

Proteriades remotula (Cockerell) (new combination)

Osmia remotula Cockerell, 1910, Can. Ent., 42: 170.
Erythrosmia remotula. Bray, 1917, Pomona Jour. Ent. ZooL, 9 : 96.
Ho-plitina remotula, Cockerell, 1922, Amer. Mus. Novitates, 40: 6.
Hoplitis remotula, Sandhouse, 1939, Mem. Ent. Soc. Washington, 1: 140.

This species is not common and only a few specimens are known.
The female can easily be recognized by the bulging upper half of
the clypeus and the depressed lower half, as well as by the broadly
rounded margin. It differs from other species, except incanescens ,
by the five-segmented maxillary palpi. The male, here described

1. Seen from the side cf the body this projection appears as a tooth. Actually the middle
coxa is produced ventrally, mesad and basad of the base of the trochanter, to form a small
transverse rounded lamella.

396 The University Science Bulletin

for the first time, differs from other species in the almost semicircu-
lar emargination of the seventh metasomal tergum.

Male: Black, including abdominal sterna; first to third meta-
somal terga and sides of fourth red; posterior margins of third to
fifth terga broadly testaceous. Mandibles red subapically; distal
ends of tarsal segments reddish ; under surface of flagellum and parts
of tegulae dark brown. Wings dusky hyaline, stigma and veins
black. Head much broader than long; inner orbits distinctly con-
verging below. Genal areas over two-thirds as broad as eyes seen
from side. Distance between posterior ocelli subequal to distance
from one of them to posterior margin of vertex and to eye margin.
Clypeal margin broadly convexly rounded, slightly crenulate, an-
gulate sublaterally, margin not thickened. Mandible with outer
tooth acute, inner blunt and much shorter than outer. Maxillary
palpi five-segmented; hairs of galeae and labial palpi sparse and
scarcely hooked. Hypostomal carinae low. Antennae short, first
and last flagellar segments markedly longer than broad, second to
fifth broader than long, remaining segments about as long as broad.
Middle coxae not toothed. Posterior coxae not lamellate; carinae
very inconspicuous. Sixth metasomal tergum only feebly angulate
at sides. Sides of seventh tergum feebly angulate, not emarginate;
apex strongly bilobed, emargination between lobes semicircular,
about twice as broad as apex of a lobe. First metasomal sternum
flat, sparsely pubescent, with irregular fringe along posterior margin.
Second sternum without swellings, its posterior margin weekly emar-
ginate medially, fringe feeble. Third, fourth, and fifth sterna con-
spicuously fringed, the fringes conforming to the broadly concave
sternal margins. Sixth sternum without the usual ferruginous fold,
but with a feeble longitudinal median ridge ending in a small pro-
jection between the broad apical lobes of the sternum. Punctation
of head and thorax fine, that of vertex coarser than that of meso-
scutum, clypeal margin broadly impunctuate but not smooth and
shining. Pubescence white, pale ochraceous on dorsum of head and
thorax, largely covering face to well above antennal bases, rather
long and abundant on thorax, not forming abdominal bands.
Length, 5.2 mm. ; fore wing, 4 mm.

Female: Black, first three metasomal terga ferruginous red,
sometimes narrowly blackish at the middle. (In a Modoc County
female, third tergum black, with a red patch on each side at the
base and a black patch on middle of the base of first two terga.)
Tegulae dusky red to black, mandibles more or less reddish on

Bees of the Genus Proteriades 397

apical half, but the flagellum dark. Wings dusky hyaline, the
stigma and veins nearly black. Head as broad as thorax, the genal
areas nearly as broad as eyes seen from side. Distance between
posterior ocelli equal to distance from one of them to posterior edge
of vertex, less than distance from one of them to eye margin. Eyes
rather strongly converging below. Clypeus large, bulgingly convex
at base, apical two fifths abruptly depressed and only slightly arched
from side to side, this part strongly projecting over labrum and
bases of mandibles, its margin rounded out, although sometimes a
little truncate medially, this truncation shorter than distance from
end of truncation to eye margin, margins laterad of truncation
gently concave, entirely without sublateral angles. Maxillary palpi
five-segmented. Mandibles moderately expanded at apices and a
little constricted near the base, the two outer apical teeth acute,
the middle tooth closer to outer tooth than to inner one, the latter
subacute, short, separated from middle tooth by a rounded emargi-
nation. Head and thorax closely and finely punctured. Punctures
of clypeus a little coarser than those elsewhere, close at sides of
disk, but widely separated on middle of basal convex area and on
the apical depression. Pubescence rather thin and long, mostly
erect on head and thorax, and whitish except more or less tinged
with ochraceous on vertex and mesonotum. Hair of mesoscutum
very short and fine, with longer erect hairs interspersed, but these
mostly shorter than the hair of face and scutellum. Abdomen with
appressed hair, denser on the three apical segments, but not forming
bands. Hair on sides of first tergum longer and erect. Length, 5.5
to 6.5 mm.; fore wing, 3.8 to 4.5 mm.

Type locality: Claremont, Los Angeles County, California. Other
specimens are as follows: One female. La Crescenta, Los Angeles
County, California, on Cryptantha intermedia, May 5, 1935 (P. H.
Timberlake) ; one female, Newhall, Los Angeles County, California,
April 20, 1940 (R. M. Bohart) ; two females. Riverside, California,
on same flower, April 6, 1945 (P. H. Timberlake) ; one male, one
female, Pinacles National Monument, San Benito County, Cali-
fornia, April 24, 1948, on Cryptantha (P. D. Hurd) ; one female.
Rock City, Mount Diablo, Contra Costa County, California, May
24, 1940 (E. G. Linsley) ; one female (unusually dark in coloration),
Fandango Pass, Modoc County, California, July 10, 1946, on As-
clepias (P. D. Hurd, R. F. Smith) from the collection of Doctor U.
Lanham.

The type is in the United States National Museum.

398 The University Science Bulletin

We are indebted to Dr. E. G. Linsley and Mr. P. D. Hurd of the
University of California for the opportunity to study the male of
this species.

Proteriades incanescens (Cockerell) new combination

This is a medium sized or large species, the female (male un-
known) of which has the upper portion of the clypeus strongly con-
vex, shining, with punctures separated by more than their diameters.
In this respect it resembles reniotula and to some extent hamuli-
cornis. Both of these species are smaller and neither has the strongly
produced and clearly truncate clypeal margin which characterizes
incanescens. Another distinctive feature of incanescens is the long
antennal scape, which is almost five times as long as broad and
longer than the distance between the inner margins of the antennal
sockets. In other species the scape is usually less than four times
as broad as long and scarcely longer than the distance between the
inner margins of the antennae sockets.

Female: Black, first three metasomal terga and sometimes base
of fourth red. Mandibles dark red, infuscated basally. Antennal
fiagellum brown, paler beneath. Legs brownish black, tarsi paler,
distitarsi rather light brown. Tegulae brownish black. Wings clear,
veins and stigma blackish. Head about as wide as thorax and about
as long as broad. Genal areas as wide as eye seen from side. Dis-
tance between posterior ocelli subequal to distance from one of them
to eye margin and to posterior edge of vertex. Inner orbits con-
verging below. Upper two thirds of clypeus strongly convex, lower
third depressed, margin strongly produced over labrum to a broad
truncation which is feebly emarginate medially, angles demarking
truncation conspicuous, separated by distance equal to distance
from one of them to lateral angle of clypeus, sublateral angles of
clypeus virtually absent. Mandibles narrowed near the base by a
gentle emargination of the inner margin as usual in this group;
apices with distance between outer and middle teeth less than that
between middle and inner teeth, all teeth acute. Hypostomal car-
inae highest behind angle, gradually narrowed anteriorly. Maxil-
lary palpi five-segmented. Punctation rather coarse and sparser
than in many species, convex upper portion of clypeus coarsely
punctured, median portion shining with punctures separated by
several diameters but laterally puctures are closer; mesoscutum at
least as coarsely and less closely punctate than vertex and at least
as coarsely punctate as mesepisternum, areas mesad to parapsidal

Bees of the Genus Proteriades 399

furrows sometimes a little more finely and closely punctured than
adjacent areas. Pubescence abundant and white except for vertex
and disk of mesoscutum where it is pale ochraceous; supraclypeal
and paraocular areas densely pubescent nearly to level of anterior
ocellus; thorax rather more densely pubescent than usual; first
three metasomal terga with narrow apical bands of pubescence
which are broken medially ; fourth more densely pubescent but with
a denser apical band; fifth and sixth uniformly covered with white
pubescence. Scopa yellowish white.

Proteriades incanescens incanescens (Cockerell) new combination

Hoplitina incanescens Cockerell, 1922, Amer. Mus. Novitates, 40: 7.

This subspecies differs from the following by its larger size, the
presence of red on the base of the fourth metasomal tergum, and
perhaps by the slightly coarser punctation of the mesoscutum.
Length, 7.0 to 7.5 mm.; fore wing, 4.8 mm.

This subspecies is known from two specimens, the type from
Grand Canyon, Arizona, May 24, 1918, and another from the same
locality (7,000 feet altitude), June 5, 1940 (R. M. Bohart). The
type is in the collection of the American Museum of Natural His-
tory and was lent for study through the courtesy of Dr. Mont A.
Cazier.

Proteriades incanescens nevadensis new subspecies

This subspecies differs from the above in its smaller size, the ab-
sence of red on the base of the fourth metasomal tergum, and the
slightly finer mesoscutal punctation. Length, 6.0 to 6.5 mm.; fore
wing, 4.0 to 4.2 mm.

Holotype female: Kyle Canyon, Charleston Mountains, Nevada,
May 24, 1940 (G. E. Bohart). One female paratype: Owens Val-
ley, Mono County, California, 7,000 feet altitude. May 31, 1941,
(R. M. Bohart).

The holotype is in the collection of the California Academy of
Sciences, San Francisco, California.

A specimen from Deep Creek, San Bernardino County, California,
May 9, 1936 (E. G. Linsley) has the clypeus less clearly truncate
than in incanescens, and has the interocular and interocellar dis-
tances more as in remotula. It probably represents either a new
species or, more likely, an intergrade between nevadensis and remo-
tula. If the latter interpretation is correct, incanescens and ne-
vadensis should both be considered subspecies of remotula.

400 The University Science Bulletin

Proteriades tristis Michener

Proteriades tristis Michener, 1936, Bull. Southern Calif. Acad. Sci., 35 : 92.

This is a rather large species, black with the red confined to the
first two abdominal segments. It thus resembles the structurally
very different P. jacintana.

Male: Black, large lateral areas on the first metasomal tergum,
nearly meeting dorsally, and small areas at sides of second tergum
red. (One specimen has red areas at sides of third tergum.) Poste-
rior margins of first six terga broadly testaceous. Flagellum and
parts of tarsi dark brown. Mandibles red subapically. Tegulae
amber colored. Wings dusky, veins and stigma black. Head as
wide as thorax. Genal areas nearly as wide as eyes seen from side.
Distance between posterior ocelli less than distance from one of
them to posterior margin of vertex or to eye margin. Inner orbits
slightly converging below, except for lower portions which are
slightly divergent. Flagellar segments mostly slightly longer than
broad but first very short and broader than long. Clypeal trunca-
tion broader than labrum, delimited by distinct angles, slightly con-
cave, sometimes slightly crenulate; margin of truncation distinctly
and broadly thickened. Mandibles with outer tooth not greatly
longer than inner. Maxillary palpi four-segmented. Middle coxae
each with distinct tooth in front of base of trochanter. Posterior
coxae not lamellate, although carinae are very high. Sixth tergum
with a distinct small tooth at each side. Seventh with a conspicuous
emargination on each side separating the rounded lateral teeth from
the bilobed median projection. First metasomal sternum but little
thickened, without protuberance. Second to fifth sterna fringed,
fringe of second short and rather irregular, of third long and con-
forming to emargination of sternum, of fourth shorter and rather
sparse, of fifth still shorter but dense. Sixth sternum with the usual
ferruginous fold, the summit of which is flattened and provided with
short pubescence; apex of fold produced into a spine. Punctation
rather fine, finer and closer on clypeus and lower parts of paraocular
areas than elsewhere on head, markedly coarser on mesepisterna
than elsewhere on head and thorax. Punctation of abdomen rela-
tively coarse, many of punctures larger than those of mesoscutum.
Pubescence white, rather sparse, not completely hiding surface even
on clypeus, forming weak apical bands on lateral portions of metaso-
mal terga. Length, 5.6 to 7.0 mm.; fore wing, 4.0 to 4.8 mm.

Female: Coloration similar to male, red of second tergum often
greatly reduced or even absent. Head as wide as thorax, broader

Bees of the Genus Proteriades 401

than long. Genal areas as wide as eye seen from side. Distance
between posterior ocelli slightly less than distance from one of them
to eye margin or to posterior margin of vertex. Eyes slightly con-
verging below. Clypeus not strongly bulging, margin with distinct
truncation broader than distance from end of truncation to lateral
angle of clypeus; angles demarking ends of truncation distinct, sub-
lateral angles weak and close to ends of truncation. Mandibles nar-
rowed near base by emargination of inner margin; apex with teeth
acute. Maxillary palpi four-segmented, rather long and slender.
Punctation much as in male. Clypeus closely punctured, a little
more coarsely so than rest of head. Pubescence rather sparse, white,
faintly ochraceous on dorsum of head and thorax, forming apical
bands of white pubescence on posterior margins of first four meta-
somal terga and covering surfaces of fifth and sixth terga. Scopa
white. Length, 6.0 to 7.0 mm.; fore wing, 3.8 to 4.2 mm.

The holotype of this species, from Eagle Rock Hills, Los Angeles
County, California, is on deposit in the California Academy of Sci-
ences. Additional specimens are from La Crescenta, Los Angeles
County, California; Altadena, Los Angeles County, California;
Idyllwild, Ribbonwood, Vandevanter Flat, and Santa Rose Moun-
tain (6,400 and 7,500 feet altitude), San Jacinto Mountains, Cali-
fornia (all C. D. Michener) ; Mill Creek, San Bernardino Mountains,
California (P. H. Timberlake). The species has been collected on
Cryptantha intermedia and C. micrantha. Males have been col-
lected on dates ranging from May 2 to June 18, females May 5 to
June 19.

Proteriades seminigra new species

This is one of the smaller species of generalized structure. The
male is similar to that of tristis, but smaller, with much more red
on abdomen and the apex of the seventh metasomal tergum not
bilobed. The female agrees with hamulicornis in having the punc-
tures of the frons almost uniformly close, but differs in having less
red on the abdomen and the punctures of mesoscutum about as
close as those of the frons. The metasomal sterna are usually
largely black, while in other species they are usually reddened.

Male: Black, the first four metasomal terga red on each side.
First tergum red except for a median black line, the following seg-
ments increasingly broadly black in middle, the fourth with about
the middle half black. Apical margins of terga, except the last,
and especially of the third to sixth, broadly rufotestaceous. Venter
25—90

402 The University Science Bulletin

black. Mandibles red between middle and bases of teeth (or nearly
all red). Flagellum a little reddened beneath. Tarsal claws ferru-
ginous. Tegulae amber color. Wings dusky hyaline, the stigma and
veins nearly black. Head slightly broader than long, and broader
than thorax. Genal areas nearly as wide as eyes seen from side,
widest opposite middle of eyes, and narrowing above. Antennae
moderately long, with middle segments of flagellum nearly as long
as wide. Margin of clypeus with a broad, somewhat crenulate and
slightly concave truncation, the margin of which is thickened and
impunctate, this truncation broader than labrum and limited by
distinct angles. Mandibles with outer tooth much longer and
slender than inner tooth. Maxillary palpi short and plainly four-
segmented. Middle coxae each with a small ventral tooth in front
of base of trochanter; hind coxae not lamellate. Sixth metasomal
tergum obtusely toothed far to each side, the tergal margin basad
of this tooth conspicuously convex. Seventh tergum strongly and
broadly notched on lateral margins to produce obtuse lateral an-
gulations and the moderately broad apical process, the latter well
rounded on apical corners and only feebly emarginate medially.
First sternum evenly convex across apex, where it is only slightly
elevated above the following segment; pubescence sparse and evenly
distributed except for being somewhat denser along posterior mar-
gin. Second sternum without definite preapical swelling. Apical
fringes on second to fifth sterna long, becoming very dense in middle
of third sternum and conforming to broadly emarginate apical mar-
gin of fifth sternum. Ferruginous median fold on sixth sternum
well developed, flattened on summit, especially basally and pro-
vided with short pubescence, apex produced as a short spine. Head
and thorax finely and closely punctured, the punctures dense on
clypeus, and very close on frons and mesocutum. Abdomen minutely
and closely punctured, the punctures becoming sparser on middle of
discs of first and second terga. Pubescence white, forming the usual
dense brush on clypeus, and long and rather dense on cheeks, sides
of thorax and around margins of mesonotum. Hair of abdomen
short, fine, and appressed, becoming denser on apical segments, and
longer and erect along the lateral margins. Length, 4.5 mm.; fore
wing, 3.5 mm.

Female: Black, with red on sides of first three metasomal terga.
Mandibles red except at base on outer side. Head as broad as long,
the genal areas over three fourths as broad as eyes seen from side.
Posterior ocelli a little closer to edge of vertex than to the eyes, the

Bees of the Genus Proteriades 403

distance between them about equal to distance to vertical margin.
Antennae short and slender, the segments of flagellum, except apical
one, no longer than broad. Mandibles not much broadened to
apex, and constricted as usual before the base, the three teeth rather
short and subequal, all acute. Disk of clypeus gently convex at
base, apical third depressed. The apical margin of clypeus with
truncation feebly concave medially and slightly reflexed, a little
longer than the oblique sides, angles demarking truncation distinct,
margins about half-way between ends of truncation and margins of
eyes. Punctures of head and thorax of about the usual size but
more uniformly close than usual. Punctures of clypeus much less
than a puncture width apart, a narrow impunctate strip bordering
the apical truncation. Punctures of frons, vertex and mesonotum
almost uniformly less than a puncture width apart, those of meso-
pleura more separated. Abdomen very finely and closely punctured.
Pubescence white, moderately dense on face and sides of thorax.
Vertex and disk of mesoscutum comparatively nude, but provided
with a fine appressed pubescence interspersed with somewhat longer
erect hairs. Abdomen with sparse, very fine, short, appressed hair
and narrow apical bands formed of whiter, more pumose hair on
the first three terga, the discal hair becoming progressively denser
on the last three segments but not forming bands. Ventral scopa
short and whitish. Length, 5 mm.; fore wing, 3.8 mm.

Type material: See seminigra seminigra.

Proteriades seminigra seminigra new subspecies

This form differs from the following in having the inner orbits of
the eyes of the male slightly converging below and the distance
between the posterior ocelli of the male subequal to the distance
from one of them to the eye margin or the posterior margin of the
vertex. There is slightly more black on the dorsum of the abdomen
in both sexes than in yosemitensis, the red on the second meta-
somal tergum of the female occupying about the outer third on each
side, with slightly more red on first tergum and less on third.

Holotype male: Riverside, California, on Cryptantha intermedia,
April 15, 1938 (P. H. Timberlake). Allotype female. May 6, 1938,
and one male paratype April 21, 1938, with data otherwise the same.
One female paratype. La Crescenta, Los Angeles County, California,
on Cryptantha intermedia, April 19, 1936 (C. D. Michener). The
holotype and allotype are in the collection of the Citrus Experiment
Station.

404 The University Science Bulletin

Proteriades seminigra yosemitensis new subspecies

This form differs from typical seminigra in having the head of the
male broader, the inner orbits of the eyes of the male with upper
portions slightly converging below but lower portions slightly di-
verging below. The distance between the posterior ocelli of the male
is less than the distance from one of them to the nearest eye margin
or the posterior margin of the vertex. There is less black on the
abdomen in both sexes so that the first metasomal tergum may
be almost wholly red above in either sex and the second tergum of
the female often has only a small middorsal black area.

Holotype male, allotype female, and four male and six female
paratypes: Yosemite, California, 3,880-4,000 feet altitude, May
25 and 31, 1938, some on Cryptantha (R. M. Bohart). An addi-
tional male, not a paratype, from Palmdale, California, April 11,
1936 (G. E. and R. M. Bohart) . The holotype and allotype are in
the collection of the California Academy of Sciences, San Francisco,
California.

Three specimens from Big Pine Creek, Inyo County, California,
7,500 feet altitude, June 12 to 17, 1942 (R. M. Bohart) are tenta-
tively assigned to this subspecies although the single male has the
distance between the posterior ocelli as in typical seTninigra.

Proteriades palmarum (Cockerell) new combination

Osmia palmarum Cockerell, 1935, Pan-Pacific Ent., 11: 48 (part).

Anthocopa {Xerosmiaf) palmarum, Michener, 1943, Ann. Ent. Soc. Amer., 36: 83.

This is one of two small species of Proteriades without red on the
abdomen; it differs from the other, P. nigra, by the presence of an
impunctate line on the clypeus. It does not seem probable that its
male will prove to have the peculiar characters of P. xerophila
(Cockerell).

Female: Black, with no red on abdomen, except a reddish stain
for a short distance between the black disks and testaceous apical
margins of the first five metasomal terga. Mandibles red in apical
third or half, and flagellum more or less reddened beneath. Tegulae
clear amber color. Wings a little dusky hyaline, the stigma and
nervures brownish. Head broader than thorax, but no broader than
long. Genal areas as wide as eyes seen from side, widest at and be-
low the middle of eyes and narrowing above. Posterior ocelli about
the same distance apart as distance to eye margin and slightly
farther apart than distance to edge of vertex. Maxillary palpi short,
with four nearly equal segments. Mandibles moderately expanded
at apices and strongly constricted before bases by an emargination

Bees of the Genus Proteriades 405

of the inner margin of each. Three apical teeth of mandibles de-
creasing in size and acuteness from the outer to inner tooth, the
two inner teeth being separated by a rather shallow emargination,
inner tooth scarcely acute. Clypeus large, with the strongly pro-
duced, thin-edged anterior margin broadly truncate, truncation de-
marked by broadly rounded angles. Disk of clypeus gently convex,
rather narrowly depressed apically. Head and thorax closely and
finely punctured, the punctures of frons and mesoscutum mostly
less than a puncture width apart. Punctures of clypeus of about
the same closeness, but leaving a narrow smooth raised median line
and a transverse punctureless strip across the border of the trunca-
tion. Abdomen with minute close punctures. Pubescence rather
abundant and white, except that it is slightly tinged with ochraceous
on the vertex and disk of mesoscutum. Hair-bands of first four
metasomal terga broad, dense and white, widening at sides of first
tergum. Disks of following terga covered with appressed whitish
hair. Ventral scopa short and white. Length, about 5.5 mm. ; fore
wing, 4 mm.

The holotype, from Palm Springs, Riverside County, California,
at flowers of Cryptantha, March 24, 1933, is in the collection of the
Citrus Experiment Station at Riverside, California. Another speci-
men from the same locality, on Cryptantha harhigera, was collected
on March 30, 1945 (P. H. Timberlake). A third specimen is from
nineteen miles west of Kane Springs, California, on Cryptantha
angustijolia, March 25, 1933 (P. H. Timberlake).

Proteriades nigra new species

Osmia palmartim Cockerell, 1935, Pan-Pacific Ent., 11 : 48 (part).

This species is similar to palmarum, from which it differs in hav-
ing the clypeus more uniforrhly convex and more closely puctured
almost all over, and in the mandibular dentition and greater width
of the frons.

Female: Black, the mandibles at apex and flagellum beneath dark
red. Apical segment of each tarsus and the claws dark ferruginous.
Abdomen without red except in a narrow erythrized area between
the whitish apical depression and the black disk of the first five meta-
somal terga. Tegulae pale amber color. Wings faintly dusky, the
stigma and nervures dark brown. Head a little broader than thorax,
the facial quadrangle distinctly broader than it is in palmarum.
Genal areas about three fourths as wide as eyes, narrowed above.
Posterior ocelli about as far apart as the distance from each eye
and slightly farther apart than distance to edge of vertex. Inner

406 The University Science Bulletin

orbits of eyes nearly parallel. Clypeus evenly convex on basal part
and slightly and broadly depressed across the apex; apical margin
with truncation about equal to oblique sides, which are slightly
angulated (by the sublateral angles) rather close to the ends of
truncation; truncation limited by distinct angles which are much
less rounded than in palmarum. Mandibles moderately expanded
at apices and strongly constricted before bases by an emargination
of upper margin of each; outer and middle teeth short and rounded
(perhaps worn in type), the innermost (which is only slightly acute)
separated from middle tooth by a rectangular emargination. Maxil-
lary palpi consisting of four subequal segments ; hypostomal carinae
moderately high behind angles, gradually reduced at and in front
of angles. Head and thorax finely and closely punctured, the punc-
tures becoming distinctly sparser but no coarser on mesepisterna.
Punctures of clypeus partly longitudinally lengthened and conflu-
ent, so as to appear slightly sulcate and almost unifonnly close,
except for a narrow apical punctureless strip across the truncation
(and an ill-defined median impunctate line in paratype). Abdomen
closely and minutely punctured, the punctures becoming dense on
the sides of the terga. Pubescence white and moderately dense on
head and thorax, with the vertex and disk of mesoscutum well ex-
posed. First four metasomal terga each with a dense white apical
hair band, that on first much broadened on each side. Fifth and
sixth terga unifoimly covered with appressed white hair. Ventral
scopa short and whitish. Length, 5.8 mm.; fore wing, 4 mm.

Holotype female: Four miles east of Edom (now called Thousand
Palms), Riverside County, California, on Cryptantha angustijolia,
April 17, 1937 (P. H. Timberlake). One female paratype (a co-type
of palmarum in the Museum of the University of Colorado), Palm
Springs, Riverside County, California, on Cryptantha angustijolia,
April 2, 1927 (P. H. Timberlake). Another female paratype, eight-
een miles east of Desert Center, Riverside County, California, on
Cryptantha barbigera, April 13, 1949 (P. H. Timberlake). The
holotype is in the collection of the Citrus Experiment Station, River-
side, California.

Proteriades si7nilis new species

This species differs in the female from any of the other smaller
species, except bidenticauda, in having the punctures of the clypeus
dense enough to dull the surface. It differs from bidenticauda in
having the punctures of clypeus more round, the clypeal margin a
little notched on each side, the antennae shorter, and the punctures

Bees of the Genus Proteriades 407

of the frons finer and denser. Superficially it resembles a large
specimen of nanula, from which it differs in the clypeal punctation,
among other characters. This species may be the female of bas-
ingeri.

Female: Black, with first three metasomal terga and sides of the
fourth rather dark ferruginous red, but base of first tergum, a small
spot at base of second, and middle of the third, black. Mandibles
black on basal half and at apex, and red preapically. Antennae
entirely dark. Tarsi slightly brownish. Tegulae amber color.
Wings dusky hyaline, the stigma and nervures dark brown, with
subcosta nearly black. Head about as broad as long, and broader
than thorax. Inner orbits of eyes slightly converging below. Dis-
tance between posterior ocelli subequal to distance from one of them
to eye margin, slightly more than distance to posterior edge of ver-
tex. Mandibles moderately broad at apices, the teeth acute and de-
creasing in size from outermost to innermost. Maxillary palpi
short but distinctly four-segmented. Hypostomal carinae mod-
erately high posteriorly, gradually reduced toward angles and much
reduced in front of angles. Antennae short, the segments of flagel-
lum, except the last, no longer than thick. Disk of clypeus gently
convex basally, and depressed on apical fourth. Margin of clypeus
broadly rounded out medially, rather strongly but obtusely angulate
by the sublateral angles on each side, and minutely emarginate on
each side, just within the sublateral angulation, that is, between the
sublateral angles and the angles demarking the truncation proper,
the latter very feeble and recognizable largely because the slightly
thickened and shining margin of the truncation proper ends at these
angles. Head and thorax closely and rather finely punctured.
Punctures of clypeus and supraclypeal area dense, imparting a dull
effect, those of frons becoming a little separated on the convexities
in front of the posterior ocelli. Punctures of mesoscutum mostly
less than a puncture-width apart, but becoming more widely spaced
just mesad of the parapsidal lines. Punctures of mesepisterna two
to three puncture-widths apart, except on anterior margin where
they are close. Abdomen minutely and closely punctured, the punc-
tures finer and sparser on middle of disk of first tergum. Pubes-
cence white and moderately dense on face and sides of thorax, and
sparse, short and pale ochraceous on vertex and mesonotum. Hair of
abdomen whitish and thin, a little denser on apical segments, and
forming very narrow, thin, apical bands only on each side of first
three metasomal terga. Ventral scopa short and whitish. Length,
4.6 mm. ; fore wing, 3 mm.

408 The University Science Bulletin

Holotype jemale: Riverside, California, on Cryptantha inter-
media, April 7, 1938 (P. H. Timberlake), in the collection of the
Citrus Experiment Station, Riverside, California.

Proteriades nanida new species

This is distinguishable from other species, except pygmaea, by its
small size and by having the margin of clypeus of the female more
completely rounded out. The almost equally small P. similis differs
by having the clypeus very densely and finely punctured. The
males of nanula and pygmaea have the lateral margins of the seventh
tergite feebly angulated, with only a slight emargination before the
apical truncation. P. nanula differs from pygmaea by being larger
and less hairy and by having the wings duskier. Certain other dif-
ferences are indicated in the key.

Male: Black, with the abdomen, including venter, ferruginous
(sometimes almost entirely so), basal middle of the terga more or
less blackened, generally most extensively so on fourth and fifth
terga. Mandibles with a red subapical band (or almost entirely
rufous). Labrum dark testaceous. Flagellum and tarsi more or less
brownish. Tegulae dark amber color. Wings dusky hyaline, the
stigma and veins dark brown. Head as broad as long, the inner
orbits converging below a little more distinctly than in most other
species. Genal areas more than half as wide as eyes seen from side.
Clypeal truncation slightly broader than labrum, limited by distinct
angles, its margin impunctate, not thickened, slightly crenulate.
Mandibles with teeth acute, with the outer one considerably longer
than the inner tooth. Maxillary palpi tapering, four-segmented (or
sometimes with last two segments fused, thus three-segmented).
Hypostomal carina low. Distance between posterior ocelli subequal
to distance from either to eye margin and greater than distance to
edge of vertex. Antennae short, the scape short and a little swollen,
middle segments of flagellum broader than long. Middle coxae not
toothed ; posterior coxae not lamellate. Tooth on each side of sixth
metasomal tergum short and obtuse. Sides of seventh tergum shal-
lowly emarginate before the apex, so that the angulations are very
low and obtuse. Apex moderately wide, bilobed because of a median
indentation (or truncate because of absence of this indentation).
First sternum sparsely pubescent, evenly convex across the apex,
slightly elevated medially and rather abruptly declivous behind.
Second sternum planate, without preapical swelling, its apical fringe
poorly developed. Fringes on third to jfifth sterna rather long and

Bees of the Genus Proteriades 409

dense, conforming to the curvature of margins. Sixth sternum with
the usual ferruginous median fold. Puncturation of head and thorax
fine and very close, that of mesopleura much sparser than elsewhere.
Punctures of clypeus sparse apically, leaving a more or less evident
narrow impunctate space across margin. Pubescence white, rather
sparse, with frons, vertex, disc of mesoscutum and base of scutellum
well exposed. Brush of hair on clypeus and face below antennae
moderately dense. Hair of abdomen sparse, not forming bands.
Length, 3.0 to 4.0 mm. ; fore wing, 2.4 to 2.8 mm.

Female: Black, with the first three metasomal terga and sides of
fourth and fifth ferruginous red, a blackish stain or spot, however,
often more or less developed at middle of base of the basal seg-
ments. Mandibles clear red in apical half (or more), teeth black.
Flagellum and tarsi slightly reddened. Inner orbits of eyes con-
verging below. Mandibles constricted near the base and moder-
ately wide at apex, the teeth all acute, nearly equally spaced, with
the outermost one the largest. Distance between posterior ocelli
subequal to distance to edge of vertex and to distance to eye margin.
Clypeal margin rounded, angles limiting truncation virtually un-
recognizable, sublateral angles distinct and midway between ends
of truncation and lateral angles of clypeus. Head and thorax closely
punctured, the punctures rather coarse in relation to the small size
of the insect (the punctures almost as large as those of the much
larger P. semirubra) . Punctures of clypeus and mesopleura a little
sparser than elsewhere. Punctures of abdomen nearly as strong and
but slightly sparser than those of mesonotum, nearly uniform in
distribution, except that they become considerably sparser and finer
on middle of the first tergum. Pubescence moderately dense and
whitish on face, cheeks, sides of thorax and around margins of
mesonotum. First four terga each with a narrow white apical hair-
band, which is seemingly easily worn or rubbed. Last two terga
evenly covered with appressed whitish hair, partially concealing the
surface. Length, 3.5 to nearly 5 mm.; fore wing, 2.3 to 3 mm.

Type material: See P. nanula nanula.

Proteriades nanula nanula new subspecies

This is a small subspecies with the wing length not exceeding 2.8
mm., with punctation rather coarse, with the red of the abdomen
extensive so that the black areas of the first three metasomal terga
of the female do not reach the posterior margins of the terga and
occupy only one fourth or less of the width of the terga.

This subspecies occurs in the lowlands of Southern California.

410 The University Science Bulletin

Holotype male: Riverside, California, on Cryptantha intermedia,
April 3, 1938 (P. H. Timberlake). Allotype female with the same
data, but collected April 26, 1937. Seven male and nine female
paratypes from the same locality and flower, males taken from
March 12 to May 6, females from March 12 to May 9; two female
paratypes, one and one half miles west of Ferris, California, April
27 and May 4, 1938 ; two female paratypes, the Gavilan, California,
April 16, 1939, and April 17, 1938; one female paratype, Claremont,
California, May 4, 1945 (all collected by Timberlake),

The holotype and allotype are in the collection of the Citrus Ex-
periment Station.

Proteriades nanula sparsa new subspecies

Somewhat larger than typical nanula, wing length of the female
usually 3 mm., punctation finer than in nanula, black areas of ab-
domen more extensive, the black of the first three metasomal terga
reaching back to the rufous margins of the terga and on the third
tergum occupying almost the dorsal third of the tergum.

This subspecies ranges across central California from near the
coast to the east side of the Sierra Nevada, and, to judge by a single
specimen, ranges into southern California in the mountains.

Holotype male, allotype female, and two female paratypes:
Hastings Natural History Reservation, near Jamesburg, Santa Lucia
Mountains, Monterey County, California, 1,900-2,700 feet alti-
tude, on Cryptantha, June 6, 1938 (C. D. Michener). One male
paratype, some data, but June 4, 1938; two female paratypes, near
Coalinga, California, June 8, 1941 (R. M. Bohart) ; one female para-
type, Yosemite, California, 3,880 to 4,000 feet altitude, May 25,
1938 (D. W. Hendrickson) ; three male and one female paratypes,
same locality. May 25 and 31, 1938, on Cryptantha (R. M. Bohart) ;
one female paratype. Badger, Tulare County, California, on Cryp-
tantha flaccida, June 26, 1929 (P. H. Timberlake) ; two male, five
female paratypes, Lone Pine Canyon, Inyo County, California, 8,000
feet altitude, on Cryptantha, June 11, 1937 (C. D. Michener) ; four
female paratypes, Big Pine Creek, Inyo County, California, June
12, 1942 (R. M. Bohart) ; one male paratype, north fork of Bishop
Creek, Inyo County, California, 8,500 feet altitude, June 22, 1937
(C. D. Michener) . One female, Mill Creek, San Bernardino Moun-
tains, 6,000 feet altitude, on Cryptantha micrantha var. lepida, July
2, 1944.

The holotype and allotype are in the Snow Entomological Col-
lections.

Bees of the Genus Proteriades 411

Proteriades pygmaea new species

This species is similar to nanula but is slightly smaller, with
abundant white pubescence, broad white hair-bands on abdomen
and clearer wings. In both sexes the head is declivous closer behind
the ocelli than in nanula so that the posterior ocelli are closer to the
posterior margin of the vertex.

Male: Black, the abdomen ferruginous red, with a dark basal
spot at middle of second and third terga and a larger infuscation
on middle of fourth and fifth terga ; sixth infuscated basally ; seventh
entirely infuscated. Apical margins of terga, especially of the sixth
and seventh, testaceous. Mandibles testaceous red, a little dusky
at base, and clearer red at apex, the teeth piceous. Flagellum and
tarsi reddish brown, the claws ferruginous. Tibial spurs yellowish
white. Tegulae pale amber color. Wings nearly clear hyaline, the
stigma and nervures brownish. Head as long as wide, the inner
orbits slightly converging below. Genal areas more than half as
wide as eyes, seen from side. Clypeal truncation broader than
labrum, margin impunctate, crenulate, not thickened. Outer tooth
of mandibles much longer and more acute than inner tooth. Dis-
tance from posterior ocelli to nearest eye margin somewhat less than
distance between posterior ocelli; distance from posterior ocelli to
edge of vertex still less, being about as great as ocellar diameter.
Antennae short, middle segments of flagellum as long as thick. Mid-
dle coxae not toothed ; posterior coxae not lamellate. Sixth tergum
with a small obtuse tooth on each side. Seventh tergite shallowly
emarginate on each side to form the apical process, lateral angula-
tion broadly rounded; apical process moderately wide and only
slightly notched medially, so that the margin is scarcely bilobed.
Venter as in nanula except that the apical fringe is well developed
on second sternum as well as on third to fifth sterna. Head and
thorax closely punctured, the punctures rather coarse in relation
to the small size of the insect. Punctures of clypeus and frons one
puncture-width, or less, apart, and those on vertex, especially be-
tween and behind the ocelli, becoming more separated. Punctures
of mesoscutum mostly somewhat more than a puncture-width apart,
and more widely separated just mesad of parapsidal lines. Punc-
tures of mesopleura about as close as those of mesoscutum. Ab-
domen minutely and closely punctured. Pubescence white, long
and dense on face, cheeks, sides of thorax, margins of mesonotum,
posterior sides of front and middle femora and on posterior margins
of hind tibiae. First four metasomal terga each with a white apical

412 The University Science Bulletin

hair-band, but these bands much less broad and dense than in fe-
male (and easily worn so that they are usually missing). Apical
terga covered with moderately dense, appressed, white hair. Length,
3 to 3.5 mm.; fore wing, 2.6 to 2.8 mm.

Female: Similar to male. Black, the first three metasomal terga
ferruginous red, with base of first and middle of second and third
more or less clouded with blackish. Apical margin of fourth and
fifth terga rufo-testaceous. Tarsi generally more blackish than in
male. Venter more or less reddish, but suffused with black on api-
cal segments. Mandibles moderately expanded at apex, inner mar-
gin emarginate near base; the three teeth almost equally spaced,
acute, but decreasing in size from outer to inner tooth. Antennae
short, the middle segments of flagellum as long as thick. Genal
areas about two thirds as wide as eyes, seen from side. Distance
between posterior ocelli greater than distance to eye margin or to
edge of vertex. Clypeus slightly and evenly convex at base and
depressed apically, the apical margin obtusely angulate at sides and
subtruncate or broadly rounded medially, the whole having the
appearance of being strongly rounded out; angles limiting ends of
truncation more distinct than in nanula, distance between these
angles and sublateral angles much less than distance from latter to
eye margins; clypeal margin between sublateral angles and ends of
truncation slightly concave. Maxillary palpi short, with four nearly
equal, closely joined segments. Pubescence white, unusually dense
and mostly subappressed, that on the face extending more than half-
way upward between insertion of antennae and the ocelli, and that
on apex of scutellum more erect than elsewhere. Dense hair covers
the face, genal areas, margins of mesonotum, sides of thorax, pos-
terior surfaces of front and middle femora, anterior surfaces of hind
coxae and femora, and outer sides of hind tibiae. First three meta-
somal terga each with a broad white apical hair-band, broadening
toward each side on first tergum. Three apical terga covered with
rather dense appressed white hair, becoming a little denser and
somewhat bandlike at apices of fourth and fifth terga. Ventral
scopa short and whitish. Length, 3.25 to 4.0 mm.; fore wing, 2.5 to
2.8 mm.

Holotype male, allotype female and one male and two female
paratypes. La Quinta, west of Indio, Riverside County, California,
on Cryptantha barbigera, March 4, 1936 (P. H. Timberlake). One
female paratype. Palm Springs, Riverside County, California, on
same flower, March 30, 1945 (P. H. Timberlake) ; one female para-

Bees of the Genus ProterIades 413

type, seven and one half miles south of Twenty-nine Palms, River-
side County, California, on Nama demissum, May 7, 1948 (P. H.
Timberlake) ; one male, four female paratypes, two miles south of
Cathedral City, Riverside County, California, on Cryptantha, April
10, 1936 (C. D. Michener) ; one female paratype, five miles east of
Cathedral City, on Cryptantha, April 10, 1936 (E. G. Linsley) ;
and one male paratype, Panamint Springs, Inyo County, California,
on Cryptantha, April 7, 1938 (E. G. Linsley).

The holotype and allotype are in the collection of the Citrus Ex-
periment Station, Riverside, California.

Proteriades basingeri new species

This species in the male differs from others, except P. boharti, in
having a low rounded protuberance at the apex of the first meta-
somal sternum. It differs from boharti in having shorter antennae
and a shining punctureless border at the apex of the clypeus, as well
as by lacking lamellae on the posterior coxae. It is quite possible
that this is the male of similis, but basingeri has never been col-
lected with that species, although they are from the same locality.
Circumstances suggesting the association of the two as sexes of one
species are their small size, and the fact that each is known only
from Riverside where both are very rare.

Male: Black, the abdomen ferruginous, with last two terga and
middle of preceding two or three segments more or less suffused with
fuscous. Sterna almost entirely ferruginous. Mandibles reddened
on apical half, teeth blackish, flagellum and tarsi brownish, the
claws ferruginous. Tegulae dark amber color. Wings dusky hy-
aline, the stigma and veins fuscous. Head as broad as long, the
genal areas about two thirds as broad as the eyes. Inner orbits
slightly converging below. Antennae short, the scape slender and
short, the middle segments of flagellum about as long as broad.
Clypeal truncation broader than width of labrum, the margin
slightly thickened, impunctate, weakly crenulate. Teeth of man-
dibles acute, the outer one somewhat longer than inner. Distance
between posterior ocelli a little less than distance from one of them
to edge of vertex or to eye margin. Middle coxae each with blunt
ventral tooth in front of base of trochanter. Posterior coxae not
lamellate. Sixth tergum distinctly angulate at sides. Lateral mar-
gins of seventh tergum strongly angulated before apex, apex broad
and bilobed. First sternum of metasoma with a medio-apical low
rounded proturbance. Second sternum with a weak transversely

414 The University Science Bulletin

arcuate subapical swelling, its apical fringe rather thin. Fringes
of third and fourth sterna long and dense, that of fifth shorter and
very dense, all of these conforming to the broadly emarginate apical
margins of the sterna; sixth sternum with the usual longitudinal
median ferruginous fold. Head and thorax finely and closely punc-
tured, the punctures sparser and a little coarser on the mesopleura
than elsewhere. Punctures of clypeus minute and dense, but leaving
a shining, nearly punctureless apical border. Abdomen minutely
and closely punctured. Pubescence whitish, or sometimes brownish
ochreous on middle of face and on the mesonotum, dense on clypeus
and face, moderately dense elsewhere on head and thorax, except
on upper part of supraclypeal area, vertex and disk of mesoscutum,
which are well exposed; hair of these parts very short, with a few
longer erect hairs interspersed. Hair of abdomen fine and short,
not forming apical hair-bands, but denser on the apical segments
and becoming much longer along the lateral margins. Length, 4.6
to 5.5 mm.; fore wing, 3.2 to 3.8 mm.

Holotype male: Riverside, California, April 11, 1933 (A. J.
Basinger). One male paratype, same locality, April 4, 1939, on
ground (P. H. Timberlake) ; one male paratype, same locality, on
Cryptantha intermedia, March 20, 1929 (P. H. Timberlake).

The holotype is in the collection of the Citrus Experiment Station,
Riverside, California.

Proteriades reducta new species

The male of reducta is distinguishable from similar middle-sized
species by having the middle of the first sternum of abdomen
covered with conspicuous white hair and by lacking the ventral
tooth of the middle coxa which is present in allied species. The fe-
male differs from truicauda in the slightly smaller size and by hav-
ing the first three terga entirely red and clypeal margin more trun-
cate medially. From bidenticauda it differs in having the maxillary-
palpi shorter, with only two or three indistinct segments, and the
clypeal margin without a notch just within the lateral angulations.
These three species differ from hamulicornis and seminigra in having
the frons more shining and less closely punctured.

Male: Black, the first four metasomal terga ferruginous red, ex-
cept base of first and sometimes a median spot or stain on the others.
Fifth tergum (sometimes) stained with red, its apical margin and
that of fourth and sixth terga broadly testaceous. Mandibles
largely red but more or less dark at extreme bases and apices.

Bees of the Genus Proteriades 415

Flagellum and tarsi dark brown. Tegulae amber color. Wings
dusky hyaline, the stigma and veins dark brown. Head about as
broad as thorax, genal areas about three fourths as broad as eyes,
the inner orbits sHghtly converging below. Posterior ocelli a little
closer together than the distance to eye margin or to edge of vertex.
Clypeal truncation broad, very gently concave, not crenulate, not
thickened, slightly broader than labrum, demarked by distinct
angles. Mandibular teeth acute, outer distinctly longer and sharper
than inner. Maxillary palpi short with two or three indistinct seg-
ments. Antennae short, the middle segments of flagellum as long as
thick, the scape only slightly swollen. Middle coxae not toothed,
posterior coxae not lamellate. Sixth metasomal tergum with a small
distinct tooth on each side. Lateral margins of seventh tergum
strongly and broadly incised before the apex to form a strong but
blunt angulation. Apical process moderately wide and indented
medially to form two rounded lobes. First sternum with a trian-
gular medio-apical area somewhat protuberant and densely covered
with rather long white hair, which also fringes the margin. Apical
fringe of second to fourth sterna long and rather dense, fifth broadly
emarginate at apex, its apical fringe short, sixth also provided with
a short apical fringe, as well as the usual median rufo-testaceous fold
which is sharp only posteriorly and broad and covered with short
hairs anteriorly. Head and thorax finely and very closely punctured,
the punctures becoming more separated on the mesopleura. Punc-
tures of abdomen minute and close. Pubescence white, moderately
dense, becoming denser on clypeus and face nearly to level of ocelli.
Clypeal brush shortening anteriorly and anterior margin of clypeus
narrowly exposed. Pubescence of abdomen thin, short and whitish,
forming only traces of bands on the basal segments. Length, 5.0 to
5.5 mm.; fore wing, 3.8 to 4.0 mm.

Female: Black, the first three metasomal terga and sides of the
fourth more or less broadly, ferruginous red, fourth and fifth terga
with posterior margins broadly rufo-testaceous. Mandibles red
subapically. Tegulae translucent brown. Head as wide as thorax,
inner orbits slightly converging below, genal areas about three
fourths as wide as eyes seen from side. Distance between posterior
ocelli subequal to distance from one of them to eye margin, slightly
greater than distance to posterior edge of vertex. Mandibles con-
stricted near base and moderately expanded at apex, the teeth acute,
with the innermost one much shorter and a little more distant from
the middle tooth than the latter is from the outer tooth. Disk of

416 The University Science Bulletin

clypeus moderately convex basally and siibdepressed toward the
apex, apical margin broadly truncate in middle, the truncation
slightly longer than the oblique sides, not thickened, scarcely im-
punctate, demarked by distinct angles; sublateral angles closer to
the ends of the truncation than to margin of eyes, clypeal margin
between sublateral angles and ends of truncation slightly concave.
Puncturation of head and thorax fine and close but less dense than
in the male, or than in seminigra and hamulicornis. Pubescence
white and moderately developed, the vertex and disk of mesoscutum
well exposed. Abdomen with thin, short, appressed whitish hair,
becoming a little denser on apical segments, and forming narrow
apical bands at sides of the first three terga. Scopa yellowish white.
Characters otherwise agreeing with male. Length, 4.5 to 5.0 mm.;
fore wing, 3.2 to 3.4 mm.

Holotype male: Riverside, California, on Cryptantha intermedia,
May 15, 1935 (P. H. Timberlake). Allotype female, April 15, 1939,
data otherwise the same. Two paratypes with same data except
that one (male) was collected May 15, 1926; the other (female),
April 26, 1926. One paratype 1.5 miles west of Perris, Riverside
County, California, May 14, 1946 (P. H. Timberlake).

The holotype and allotype are in the collection of the Citrus Ex-
periment Station, Riverside, California.

Proteriades cryptanthae new species

This species is nearly of the same size and color as semirubra.
It is easily distinguished in the male from all other species except
caudex by having a large hornlike process on the apical middle of
the first metasomal sternum. The male differs from caudex in the
larger size and in having the ventral process thin-edged and trun-
cate at apex. From semirubra the female differs in having the hair-
bands of abdomen little developed, and the clypeal margin without
fingerlike processes.

Male: Black, the first four terga of abdomen red; fourth tergum
(often) with a large median infuscation or subapical cross band
(third tergum more rarely with a dark cross band). Apical margins
of fifth and sixth terga rufo-testaceous. Apical segments of tarsi
rufous (or brown). Mandibles with a subapical dark red blotch on
band. Flagellum moderately reddened beneath. Tegulae fuscous,
slightly reddened. Wings strongly dusky, stigma and veins piceous.
Head as broad as thorax, broader than long; genal areas two thirds
as wide as eyes seen from side. Distance between posterior ocelli

Bees of the Genus Proteriades 417

less than distance from one of them to eye margin or to posterior
margin of vertex. Clypeal margin with truncation about as broad
as labrum, and defined by distinct angles, the margin of truncation
slightly thickened, impunctate, and feebly produced medially and
sublaterally. Maxillary palpi four-segmented. Antennae shorter
than in semirubra, the middle segments of flagellum scarcely longer
than thick. Middle coxae each with a slight ventral swelling but
without a tooth; posterior coxae not lamellate, but carinae very
strong. Sixth tergum with a tooth on each side, seventh with a
larger and sharper tooth on each side, beyond which is a broad
emargination, and with the median lobe moderately wide and trun-
cate at apex, with only a trace of a median emargination. (In
semirubra the median lobe is considerably broader and distinctly
emarginate, and the lateral teeth are smaller and much closer to
apex of median lobe.) First metasomal sternum with a large ver-
tically descending process from its apex, this process convex in
front, flat behind, and thin-edged and truncate at apex. Second,
third, and fifth sterna each with apical fringe, that on third be-
coming denser at the middle, those on second and third long, and
that on fifth rather short, very dense, and conforming to the broadly
emarginate margin of the segment; fourth sternum with only a
sparse fringe. Head and thorax finely and very closely punctured,
the punctures becoming slightly more separated on scutellum, and
larger and more separated on mesopleura. Abdomen minutely and
closely punctured, the punctures less crowded toward the base.
Pubescence white, moderately dense on head, thorax and legs, and
leaving vertex and disk of mesoscutum well exposed. Hair of face
sometimes tinged with pale brown, and covering surface nearly to
level of ocelli, the clypeal beard, however, less dense than in se7ni-
rubra. Abdomen with fine short hair, most dense on apical mar-
gins and forming weak bands laterally on first four metasomal
terga. Length, 6.0 to 7.0 mm.; fore wing, 4.2 to 5.0 mm.

Female: Black, with first three metasomal terga, base of fourth
and apical margins of fourth and fifth red ; sterna black. Mandibles
red apically and along the inner margin. Coloration otherwise as
in male. Head large, as broad as thorax, genal areas considerably
broader than eyes seen from side and broadest just above middle
of eyes. Mandibles not much narrowed toward bases, inner mar-
gins a little emarginate near bases, much less broad at apices than in
semirubra, with the three teeth acute, the outer one a little longer
and more acute than the others. Clypeus rather strongly convex,

26—90

418 The University Science Bulletin

its apical margin somewhat depressed; truncation broad, demarked
by distinct angles which are (usually) farther apart than distance
from one of them to eye margin; margin of truncation thick, crenu-
late with about four to six small blunt teeth, and projecting but
little over the base of labrum. Distance between posterior ocelli
less than distance from one of them to nearest eye margin and sub-
equal to distance to edge of vertex. Puncturation nearly as in the
male. Pubescence whitish, much less dense on the face than in the
male. Ventral scopa short, tinged with ochraceous. Length, 6.4 to
7.5 mm.; fore wing, 4.0 to 4.7 mm.

Holotype male: Riverside, California, on Cryptantha intermedia,
April 18, 1928 (P. H. Timberlake). Allotype female: same data
except collected April 23, 1928. Nine male and eight female para-
types, same data except males taken March 21 to May 1, females
April 20 to June 17; one female paratype, the Gavilan, Riverside,
County, California, April 16, 1939; one female paratype, six miles
east of Temecula, California, May 9, 1936; two female paratypes
from Claremont, Los Angeles County, California, May 4, 1945; all
on flowers of Cryptantha intermedia except one rescued from a
spider and all collected by P. H. Timberlake; one additional female
paratype from Newhall, Los Angeles County, California, April 20,
1941 (R. M. Bohart, in the Bohart collection).

The holotype and allotype are in the collection of the Citrus Ex-
periment Station, Riverside, California.

Proteriades caudex new species

In the male this species resembles the considerably larger cryptan-
thae by having a very large protuberance on the first sternum; it
differs from cryptanthae by having this sternal process thick and
stumplike instead of flattened antero-posteriorly and thin-edged at
apex. The posterior coxae are lamellate, a character not shared by
cryptanthae. The female is far less distinctive but can be recog-
nized by the narrow truncation of the clypeus, on each side of which
is a distinct concavity between the end of the truncation and the
sublateral angle.

Male: Black, first three abdominal terga red, the basal concavity
of the first infuscated, the second with an infuscated dorsal patch,
the third with a rather large (or sometimes small) dorsal black patch,
the fourth broadly red laterally. Third to sixth terga with posterior
margins testaceous. Mandibles reddish subapically (or largely red),
Flagellum brown, paler beneath. Tegulae dark amber color. Wings

Bees of the Genus Proteriades 419

slightly dusky, veins and stigma blackish. Head as broad as thorax,
broader than long. Genal areas about two thirds as broad as eyes
seen from the side. Distance between posterior ocelli slightly less
than (to slightly more than) distance from one of them to eye mar-
gin and to posterior edge of vertex. Inner orbits slightly converging
below. Antennae long, scape slightly swollen, flagellar segments
all longer than broad, middle ones nearly twice as long as broad.
Clypeal truncation wider than labrum, demarked by distinct singles,
margin of truncation impunctate but not thickened, slightly crenulate.
Mandibles with teeth acute, inner one but little shorter than outer.
Middle coxae not toothed. Posterior coxae each with carina grad-
ually expanded posteriorly to form a low lamella, the posterior end
of which is produced to a rounded apex slightly beyond the end of
the rest of coxa. Sixth metasomal tergum with a small acute angle
on each side. Seventh with a large bluntly rounded basal angle on
each side, distad to which margins of tergum are deeply emarginate,
apical produced portion of tergum with conspicuous median emargi-
nation somewhat broader than the resulting lobes. First sternum
with a very large median apical protuberance with a rounded apex.
Second to fifth sterna with apical fringes, those of second to fourth
very long, conforming to the emarginate posterior edges of the
sterna. Fifth sternum with the usual ferruginous fold. Punctation
rather fine and dense, coarser and sparser on mesepisterna. Pubes-
cence white, ochraceous on dorsum of head and thorax, dense on face
nearly to level of anterior ocellus, forming only very feeble abdom-
inal bands on first five metasomal terga. Length, 4.5 to 5.8 mm.;
fore wing, 3.4 to 3.8 mm.

Female: Black, first metasomal tergum red except black area in
basal concavity and posterior median black (or infuscated) area
(sometimes connected to black area of concavity) ; second and third
terga with lateral thirds or slightly more red (fourth often red at
sides basally, occasionally with red extending to apex at extreme
sides). Coloration otherwise as in male. Head as broad as thorax,
as broad as long. Genal areas over two thirds as broad as eyes
seen from side. Distance between posterior ocelli subequal to dis-
tance from one of them to posterior edge of vertex, (usually) slightly
less than distance to eye margin. Inner orbits converging below.
Clypeus uniformly gently convex, upper part not noticeably bulging,
marginal truncation only about two thirds as long as distance from
one of the angles limiting it to lateral angle of clypeus, sublateral
angles nearer to angles limiting truncation than to lateral angles,

420 The University Science Bulletin

margin between sublateral angles and angles limiting truncation
conspicuously concave. Mandibles narrowed near bases by the
usual emargination in inner margin of each, teeth equidistant, all
acute. Maxillary palpi three- or obscurely four-segmented. Pimc-
tation fine, that of clypeus coarser than that of rest of head, about
as coarse as that of posterior part of mesoscutum, portions of lat-
ter mesad of parapsidal furrows not noticeably more coarsely punc-
tate than rest of scutum; mesepisterna conspicuously more coarsely
punctate than mesoscutum. Pubescence as in male but not so dense
on face, sparsely covering fifth and sixth terga, bands on first four
terga more conspicuous; scopa white. Length, 4.6 to 5.5 mm.; fore
wing, 3.0 to 3.4 mm.

Holotype male: Idyllwild, San Jacinto Mountains, California,
May 23, 1940 (C. D. Michener). Allotype female: Pine Flat, near
Idyllwild, San Jacinto Mountains, California, on Cryptantha mi-
crantha, June 15, 1940 (C. D. Michener). Five male paratypes,
same data as holotype but collected May 26, 1940; one male para-
type, Herkey Creek; San Jacinto Mountains, California, June 4,
1940 (C. D. Michener) . One female and four male paratypes. Keen
Camp, San Jacinto Mountains, California, on Cryptantha, May 16,
1939 (E. G. Linsley) ; seven female paratypes, same data as allotype;
one female paratype, Idyllwild, San Jacinto Mountains, California,
June 3, 1939, on Cryptantha lepida (P. H. Timberlake) ; two female
paratypes, Santa Rosa Mountain, San Jacinto Mountains, California,
on Cryptantha micrantha, May 31 and June 8, 1940 (C. D. Mich-
ener) ; one female paratype, San Jacinto Mountains, California, 1912
(J. C. Bridwell).

The holotype and allotype are in the Snow Entomological Collec-
tions.

Proteriades bidenticauda new species

HopUtina hesperia, Cockerell, 1926, Pan-Pac. Ent., 3: 87 (male, misidentification).

The male of this species is readily recognizable among Proteriades
species by the seventh metasomal tergum, the two lobes of which
have become two blunt teeth separated by a long straight margin.
A superficially similar configuration occurs in the unrelated species
deserticola, but in that form the teeth are separated by a concave
margin. The female is similar to several other species such as
hamidicornis, from which it differs in the closely punctured clypeus
among other characters.

Male: Black, abdomen red, basal concavity largely black,
(fourth tergum often with an infuscated spot dorsally), fifth to

Bees of the Genus Proteriades 421

seventh terga brownish or infuscated; posterior margins of fourth
to sixth terga broadly testaceous. Mandibles largely rufescent, red
subapically. Antennal flagellum brown, paler beneath. Tarsi
brown, paler distally. Tegulae light amber color. Wings scarcely
dusky, veins and stigma brownish black. Head as wide as thorax;
genal areas about two thirds as broad as eyes seen from sides; dis-
tance between posterior ocelli greater than distance from one of
them to posterior edge of vertex, much greater than that to eye
margin. Inner orbits of eyes parallel. Antennae with basal flagellar
segments broader than long, middle segments about as long as broad,
distal ones longer than broad. Clypeal truncation broader than
labrum, broadly thickened, gently concave, not crenulate (or
slightly crenulate), limited by distinct angles. Mandibles with
teeth acute, inner one not much shorter than outer. Hypostomal
carinae very low. Maxillary palpi four-segmented. Middle coxae
each with a small tooth in front of base of trochanter. Hind coxae
not lamellate. Sixth metasomal tergum with obtuse angle at each
side. Seventh tergum with lateral margins straight, extending back-
ward to form two blunt teeth widely separated by a margin which is
nearly straight. First sternum somewhat thickened, posterior mar-
gin declivous. Second, third, and fourth terga with apical fringes,
particularly long on third and fourth and conforming to the emar-
ginate edges of these two terga. Second sternum with a shining con-
vexity at each side, third with small convexities of the same sort.
Fifth sternum produced to a thin, transparent, unfringed margin.
Sixth sternum with the usual ferruginous fold, the lower edge of
which is broadened and covered with short hairs. Punctation rather
fine, particularly fine and close on clypeus except for the impunc-
tate margin; central part of mesoscutum between parapsidal lines
more coarsely and less closely punctate than rest of mesoscutum;
mesepisterna a little more coarsely punctate. Pubescence sparse,
white, covering face to level of antennal bases, not forming abdom-
inal bands. Length, 5.0 to 5.5 mm.; fore wing, 3.4 to 3.8 mm.

Female: Black, first four metasomal terga red, the first with
black in basal concavity, the third (and often the second, sometimes
first as well) with infuscated dorsal spot, fourth with black patch
dorsally and sometimes infuscated laterad of this patch ; fourth and
fifth terga with posterior margins broadly rufo-testaceous. Colora-
tion otherwise as in male. Head as broad as thorax, slightly broader
than long. Genal areas nearly as broad as eyes seen from side. Dis-
tance between posterior ocelli greater than distance from one of

422 The University Science Bulletin

them to eye margin or posterior edge of vertex. Inner orbits slightly
converging below. Clypeus with upper two thirds somewhat bulg-
ing, lower third depressed; truncation defined by rather obscure
angles, about as long as distance from end of truncation to lateral
angle of clypeus, sublateral angles slightly nearer to ends of trunca-
tion than to lateral angles, margins between ends of truncation and
sublateral angles slightly concave. Mandibles narrow near the base
because of the usual emargination of the inner margin, tridentate
(with the teeth blunt in most specimens probably because of wear).
Maxillary palpi four-segmented. Punctation fine, that of clypeus
close and but little coarser than that of rest of head. Punctures of
convexities in front of ocelli slightly sparser than those of adjacent
areas. Mesoscutum with an ill-defined zone of coarse and widely
separated punctures between parapsidal furrows, as in male; punc-
tures of mesepisterna but little coarser than those of this zone.
Pubescence sparse, white, forming only very weak apical bands at
sides of first four metasomal terga; fifth and sixth terga sparsely
covered with white pubescence. Length, 4.8 to 5.4 mm. ; fore wing,
2.8 to 3.6 mm.

Holotype male, allotype female: Mazourka Canyon, Inyo Moun-
tains, Inyo County, California, 9,500 feet altitude, June 1, 1937
(C. D. Michener). Seven male and one female paratypes, same
data; two male paratypes, same locality, 8,000 feet altitude. May
22, 1937; seven male and two female paratypes, same locality, 7,500
feet altude, May 21, 1937, one on Cryptantha; four male and one
female paratypes, same locality, 7,000 feet altitude, two on Cryp-
tantha; one male and two female paratypes. Wild Rose Canyon,
Panamint Mountains, Inyo County, California, 7,000 feet altitude.
May 27, 1937; one female paratype. Tuber Canyon, Panamint
Mountains, Inyo County, California, 8,000 feet altitude, June 18,
1937 (all C. D. Michener, collector) ; two male paratypes from the
type locality (no altitude record). May 21, 1937 (N. W. Frazier) ;
two male paratypes, Inyo Mountains, Inyo County, California,
June 1, 1937 (E. C. Van Dyke) ; one male paratype. Silver Canyon,
White Mountains, Inyo County, California, May 11, 1926 (J. 0.
Martin).

The holotype and allotype are in the Snow Entomological Col-
lections.

This species emerges in early spring in the high desert ranges
where it occurs and almost all the specimens collected were found
resting on rocks or sand before the Cryptantha flowers were open.

Bees of the Genus Proteriades 423

Proteriades truicauda new species

In this moderate sized species the male is very readily recognized
by the much produced, scoop-shaped seventh metasomal tergum
which is not at all bilobed and not angulate on the sides. The
female is markedly smaller than the male and differs from a series
of structurally rather similar species such as tristis and reducta by
having the red coloration restricted to the sides of the metasomal
terga but exending back to the fifth or sixth tergum.

Although as the specimens were collected it was felt that the
males and females represented a single species, we subsequently
doubted this because the males were so much larger than the fe-
males. However, males are larger than females in several other
species of Proteriades (e. g. xerophila, deserticola) . The males and
females here united under the name truicauda agree in having red
extending far back on the sides of the abdomen but restricted dor-
sally, in the broad translucent rufo-testaceous margins of the meta-
somal terga, in the segmentation of the maxillary palpi, and in
having the punctation a little coarser than in allied species.

Male: Black, the metasomal terga light red laterally; first three
terga with dark brown or blackish dorsal area, occupying dorsal half
of each tergum, so that the lateral quarters are red, these dark
dorsal areas narrowed anteriorly on each segment, and terminated
posteriorly by the broad translucent rufo-testaceous margin of each
segment; fourth tergum similar but with dark area extending farther
toward each side ; fifth and sixth with dark areas not narrowed an-
teriorly and nearly reaching lateral margins, the red that is present
heavily infuscated on sixth, the posterior margins broadly rufo-testa-
ceous as on preceding terga; seventh tergum wholly dark red,
heavily infuscated in certain areas. Abdominal sterna black.
Mandibles faintly reddish subapically. Antennal flagellum, tarsi,
apices of tibiae and apices of femora dark brown. Tegulae trans-
parent, pale honey color, appearing dark only because of dark
sclerites beneath them. Wings dusky hyaline, the veins and stigma
black. Head as wide as thorax, slightly broader than long. Genal
areas about two thirds as wide as eyes seen from side. Eyes some-
what swollen, inner orbits slightly converging below. Margin of
clypeal truncation slightly thickened, broadly impunctate, strongly
crenulate because of five or six small projections which are largest
medially, truncation broader than labrum, demarked by distinct
rounded angles. Distance between posterior ocelli subequal to dis-
tance from one of them to eye margin, slightly less than distance

424 The University Science Bulletin

from one of them to posterior margin of vertex. Mandibular teeth
subequal, outer tooth sharper than inner. Maxillary palpi short,
tapering, four-segmented. Antennae long, flagellar segments except
the first longer than broad. Hypostomal carinae very low. Mid-
dle coxae each with ventral angle in front of base of trochanter.
Posterior coxae not lamellate. Sixth metasomal tergum with a small
distinct tooth at each side. Seventh tergum large, much excerted
with a middorsal depressed area, under surface strongly concave.
Lateral margins straight (or nearly so), apical margin slightly con-
vex (to nearly straight). First metasomal sternum thinly haired,
the pubescence denser posteriorly; posterior margin of sternum
strongly thickened and considerably elevated above second sternum.
Second sternum large, with a strong shining convexity at each side,
fringe long, sparse medially. Third sternum with small convexities
at each side and a long fringe. Fourth sternum with posterior edge
broadly emarginate, the shorter fringe conforming to the emargina-
tion. Fifth sternum similarly emarginate, practically without a
fringe. Sixth sternum with the reddish median fold, sharp posteri-
orly, broad and covered with short hairs anteriorly. Head and thorax
rather finely punctured, clypeus and lower two thirds of supraclypeal
area very finely and closely so, vertex with punctures separated by
nearly a puncture width, much of mesoscutum with punctation sim-
ilar to that of vertex but areas mesad of parapsidal furrows with
much coarser and more widely separated punctures; mesepisterna
with even coarser and more widely separated punctures; abdomen
rather finely punctured. Pubescence (badly worn) whitish, sparse,
not completely covering surface even on clypeus, not forming ab-
dominal bands. Length, 6.5 to 7.5 mm. (abdomen strongly curled
under) ; fore wing, 4.1 mm.

Female: Black, the sides of the abdominal terga red, most
broadly so on the first tergum and decreasingly so toward apex,
so that the sixth tergum is nearly all black. Depressed apical mar-
gin of first fine terga pale testaceous. Mandibles dark red in apical
half. Antennae and tarsi dark, the claws and apices of femora and
tibiae ferruginous. Tegulae pale translucent amber. Wings dusky
hyaline, the stigma and veins blackish. Head about as broad as
long and broader than thorax. Inner orbits of eyes slightly con-
verging below. Posterior ocelli about as far apart as distance to
edge of vertex, and the distance from either to nearest eye margin
slightly greater. Disk of clypeus moderately convex and a little
depressed toward the apex, the punctures leaving a narrow smooth

Bees of the Genus Proteriades 425

space across apical margin. Margin of clypeus so weakly angu-
lated at sides and so weakly truncate in middle that it appears well
rounded out, sublateral angles very close to angles limiting end
of truncation and separated from them by a small emargination.
Mandibles constricted by an emargination of the inner margin near
the base, moderately expanded at apex, the two outer teeth subequal
and moderately acute, the innermost tooth nearly rectangular and
separated from the middle tooth by a rounded emargination. Max-
illary palpi short, tapering, four-segmented. Head and thorax
rather closely punctured, the punctures coarser and sparser on the
mesepisterna than elsewhere. Punctures on middle of mesoscutum
less than a puncture width apart, areas mesad of parapsidal lines
more coarsely and sparsely puctured. Pubescence whitish, mod-
erately dense on face, cheeks and thorax, with the vertex and disk
of mesoscutum well exposed. First to fourth terga each with a
rather broad white apical hair-band, and disks of fifth and sixth
terga thinly covered with appressed whitish hair. Length, 5.5 to
6.0 mm. ; fore wing, 4.0 mm.

Holotype male: Santa Rosa Mountain, San Jacinto Mountains,
Riverside County, California, on Cryptantha micrantha, June 8,
1940 (C. D. Michener). Allotype female, same locality, June 18,
1940 (C. D. Michener). Two male and two female paratypes, all
from the same locality and on the same flower, one of each sex May
31, 1940 (C. D. Michener) and one of each sex June 8, 1940 (P. H.
Timberlake).

The holotype and allotype are in the Snow Entomological Col-
lections.

Proteriades hamulicornis new species

The male of this species is easily recognizable by numerous char-
acters such as the very broad face with the inner orbits diverging
below, the sharply pointed last antennal segment suggestive of this
structure in Hoplitis, subgenus Alcidamea, and the spined posterior
coxae. By contrast, the female, which is apparently to be asso-
ciated with these males, is unspecialized in structure and similar
to several other species, such as bidenticauda. It differs in having
the prominences in front of the ocelli densely punctured without
shining interspaces and in the widely separated punctures of the
convex upper portion of the clypeus.

Male: Black, first three metasomal terga red except concavity of
first which is black, and longitudinal median infuscated streak on
third (and sometimes second). Fourth to sixth terga with broad

426 The Univeksity Science Bulletin

rufo-testaceous (or sometimes translucent testaceous) posterior mar-
gins, the fourth also (usually) red laterally and basally. Mandibles
red subapically and along inner margins (or very largely red). An-
tennal flagellum brown, pale brown beneath. Tegulae testaceous,
infuscated anteriorly. Legs brownish black, tarsi brown, lighter
toward apices. Wings slightly dusky, veins and stigma brownish
black. Head slightly broader than thorax, much broader than long,
inner orbits slightly diverging below, more strongly so in lower
thirds. Genal areas nearly two-thirds as wide as eyes, seen from
sides. Distance between posterior ocelli subequal to distance from
one of them to eye margin and to posterior edge of vertex. Antennae
rather long and thickened; scape swollen, pedicel unusually large;
fiagellum progressively thickened toward apex, subapical segment
broadest, most of the segments slightly longer than broad, apical
segment drawn out into a slender curved point as in Hoplitis,
subgenus Alcidamea. Clypeus very short and broad, three times
as broad as long, scarcely overhanging base of labrum; anterior
margin between lateral angles of clypeus scarcely produced; trun-
cation about as wide as labrum, limited by very weak angles,
its margin thickened and very feebly crenulate or wavy. Mandibles
narrowed at apices so that the two teeth are crowded together, outer
tooth much longer than inner but both acute. Hypostomal carinae
very low and inconspicuous except where each is elevated to a tooth
at angle. Maxillary palpi three-segmented, last segment longer
than either of the others and sometimes with a feeble constriction
indicating division into two segments. Middle coxae not toothed
although slightly swollen in front of bases of trochanters. Posterior
coxae with inner surfaces broadly concave, carinae scarcely evi-
dent; inner apical portion of each posterior coxa prolonged into a
long spine. Sixth metasomal tergum of male with a small angle at
each side. Seventh tergum with a broad emargination on each side,
an obtuse rounded angle basad of emargination; median portion of
tergum produced, notched medially forming two broad lobes. First
metasomal sternum without a protuberance, thickened, its posterior
margin abruptly declivous. Second sternum but briefly fringed;
third to fifth with long conspicuous fringes; sixth sternum with the
usual reddish median fold, the summit of which is flattened and pro-
vided with short hairs. Punctuation fine and dense, coarser on
posterior part of mesoscutum, on scutellum, and mesepisterna than
on vertex. Pubescence white, covering face nearly to level of an-
terior ocellus (or worn off above antennae) ; hypostomal areas

Bees of the Genus Peoteeiades 427

densely pubescent; abdominal bands virtually absent. Length, 4.8
to 5.2 ram. ; fore wing, 3.0 to 3.5 mm.

Female: Black, first three metasomal terga red, concavity of
first black (third sometimes with an infuscated dorsal patch).
Fourth tergum red basally (sometimes also laterally). Fourth and
fifth terga testaceous posteriorly. Anterior margin of clypeus red.
Coloration otherwise as in male. Head about as wide as thorax,
as long as broad. Genal areas about three fourths as wide as
eyes seen from side. Distance between posterior ocelli subequal to
distance from one of them to posterior edge of vertex, slightly less
than (or subequal to) distance to eye margin. Inner orbits very
slightly converging below. Clypeus convex, lower fourth depressed.
Clypeal margin with truncation rather narrow, angles delimiting it
weak, distance between these angles less than distance from one of
them to lateral angle; sublateral angles more distinct and closer
to angles demarking truncation than to lateral angles, margin on
each side of each sublateral angle gently concave. Mandibles nar-
rowed subbasally by a broad emargination of the upper margin,
as usual in the group; teeth acute, although upper one scarcely so.
Hypostomal carinae and antennae of the usual type. Head and
thorax finely punctured except for clypeus which is more coarsely
so with the punctures of the upper convex portion separated by
fully their diameters; punctures of prominences in front of ocelli
as close as those of supraclypeal area and vertex; punctures mesad
of parapsidal furrows not coarser or sparser than those of other
areas of mesoscutum; punctures of mesepisterna not coarser than
of mesoscutum. Pubescence white, dense on paraocular areas and
around antennal bases, forming feeble apical bands on sides of first
four metasomal terga, and partially covering surfaces of fifth and
sixth terga. Scopa whitish. Length, 4.0 to 5.0 mm.; fore wing,
3.0 to 3.4 mm.

Holotype male: Mazourka Canyon, Inyo Mountains, Inyo
County, California, 7,500 feet altitude. May 21, 1937 (C. D. Miche-
ner). Allotype female, same data, on Cryptantha. One male para-
type, same locality, 7,000 feet, May 20, 1937, on Cryptantha; one
male and one female paratype, same locality, 9,500 feet, June 1,
1937 (all C. D. Michener) ; one male paratype, same locality and
date but labeled merely Inyo Mountains, Inyo County, California
(E. C. Van Dyke). A small female specimen (not a paratype) is
from Wild Rose Canyon, Panamint Mountains, Inyo County, Cali-
fornia, May 27, 1937, 7,000 feet altitude, on Cryptantha (CD.
Michener) .

428 The University Science Bulletin

The holotype and allotype are in the Snow Entomological Collec-
tions.

Proteriades xerophila (Cockerell) new combination

Osmia xerophila Cockerell, 1935, Pan-Pac. Ent., 11: 45.

Anthocopa xerophila Linsley and McSwain, 1942, Amer. Midland Nat., 27 : 408.
Anthocopa ( Xerosviia) xerophila, Michener, 1943, Ann. Ent. Soc. Amer., 36 : 81 ; Michener,
1949, Jour. Kans. Ent. Soc, 22 : 54.

Male: Black, posterior margins of first six abdominal terga tes-
taceous. Antennal flagellum brown, much paler beneath than
above. Mandibles red subapically. Apices of femora, bases of
tibiae, apices of tibiae and much of tarsi ferruginous; tegulae rufo-
testaceous. Wings only faintly dusky, veins and stigma dark brown.
Head about as broad as thorax. Genal areas two thirds to three
fourths as broad as eyes seen from side. Distance between posterior
ocelli equal to or slightly less than distance from one of them to
posterior margin of vertex, equal to distance to eye margin. Inner
orbits very slightly converging below. Antennae long, all flagellar
segments markedly longer than broad. Clypeus with lower margin
depressed, margin produced to a lobe at each side which extends
farther downward than the truncation, the latter nearly as broad as
labrum, crenulate. Mandibles with outer tooth but little longer
than inner. Maxillary palpi four-segmented. Hypostomal carinae
very low. Middle coxae each with a long tooth arising in front of
base of trochanter. Sixth metasomal tergum with a small tooth at
each side. Seventh tergum with a small but distinct lateral emar-
gination on each side near the apex setting off a distinct short lat-
eral tooth; apical portion of tergum strongly tridentate, the lateral
teeth long, parallel, blunt, the median tooth much shorter and
sharply pointed; dorsal surface of this tergum largely shining and
impunctate. First sternum with a large median apical protuber-
ance, margined by a carina, posterior face of protuberance flattened.
Second sternum with an apical fringe which is interrupted medially.
Third and fourth sterna with long fringes. Fifth sternum not
fringed. Sixth sternum with the usual reddish median fold, the
summit of which is flattened; this fold shorter than usual, ter-
minating in a spine; on either side of base of fold is a projecting
lobe. Punctation very fine and dense, finer on mesoscutum than on
vertex, coarser on mesepisternum. Pubescence dense, white, obscur-
ing surface on lower part of face, forming conspicuous apical bands
on first five metasomal terga. Length, 7.2 to 8.8 mm.; fore wing,
5.0 to 6.0 mm.

Female: Coloration similar to that of male. Genal areas broader

Bees of the Genus Proteriades 429

than eyes seen from side except in very small specimens where they
are equal to eyes in width. Distance between posterior ocelli con-
spicuously less than distance from one of them to eye margin or to
posterior margin of vertex. Inner orbits slightly converging below.
Upper portion of clypeus scarcely bulging; truncation as broad as
distance from its end to lateral angle of clypeus, demarked by con-
spicuous angles; sublateral' angles nearer to ends of truncation than
to lateral angles, margins between sublateral angles and ends of
truncation concave. Mandibles broad apically, apical teeth acute,
narrowed near bases by the usual broad emargination of inner mar-
gin of each. Maxillary palpi four-segmented. Hypostomal carinae
high. Punctation fine, that of clypeus coarser than that elsewhere
on head and thorax, that of mesoscutum very slightly finer than that
of vertex, that of mesepisterna coarser than elsewhere on thorax.
Pubescence abundant and white, dense on face, forming conspicuous
bands on first four metasomal terga, scattered on fifth tergum, and
nearly covering sixth. Length, 6.5 to 8.0 mm.; fore wing, 4.5 to 6.0
mm.

The type of this species, from Palm Springs, Riverside County,
California, is in the collection of the Citrus Experiment Station at
Riverside, California. A number of other specimens have been col-
lected at the same locality, most of them on Cryptantha barbigera,
females from March 21 to April 16, males from March 22 to 30
(P. H. Timberlake). Other specimens are from Tub Canyon,
Borego, San Diego County, California, February 28, 1947, and
Borego, March 6, 1947 (Grace H. and John L. Sperry). The species
is also known from Little Lake, Inyo County, California.

Proteriades boharti new species

This species is distinguishable at once from all other known
Proteriades except caudex by the male posterior coxae, the inner
ventral carina of each of which is expanded posteriorly into a
lamella. This species differs from caudex by the absence of a large
protuberance on the first sternum.

Male: Black, the first four metasomal terga red, the fourth with
a black area middorsally, the third with an infuscated middorsal
longitudinal streak (or with an infuscated patch) ; fifth tergum red
at extreme sides; fifth and sixth terga with posterior margins rufo-
testaceous; sterna blackish. Mandibles red subapically, teeth
piceous. Antennal flagellum brown, paler beneath. Legs brownish,
black. Head as wide as thorax, wider than long. Genal areas about

430 The University Science Bulletin

two thirds as broad as eyes seen from side. Distance between pos-
terior ocelli equal to distance from one of them to eye margin and to
posterior edge of vertex. Inner orbits slightly converging below.
Antennal scape swollen; flagellar segments longer than broad.
Clypeal margin with truncation broader than labrum, very slightly
concave, slightly thickened, weakly crenulate, broadly impunctate.
Mandibles with inner tooth not greatly shorter than outer, both
acute. Hypostomal carinae very low, almost absent in front of
angles. Maxillary palpi apparently four-segmented. Middle coxae
not toothed but swollen in front of bases of trochanters. Posterior
coxae with carinae elevated posteriorly to form a strong lamella
on each coxa, each lamella abruptly truncated at its posterior end.
Sixth metasomal tergum with a small but distinct angle at each
side. Seventh tergum with broad lateral emargination on each side
separating basal angle from the median apical portion of the tergum
which is conspicuously bilobed by reason of a median emargination.
First metasomal sternum with a rounded subapical protuberance.
Second to fom'th sterna with fringes of long hairs conforming to the
broadly emarginate posterior margins. Fifth sternum with the usual
reddish fold, the summit of which is flattened and provided with
short pubescence, the posterior end of which is produced into a short
spine. Punctation fine and dense, that of clypeus finer and denser
than rest of head, mesepisterna more coarsely and less closely punc-
tured than mesoscutum, especially below. Pubescence white, ochra-
ceous on vertex and dorsum of thorax, forming feeble abdominal
bands which appear to be easily worn off. Length, 4.0 to 5.0 mm.;
fore wing, 2.8 to 3.5 mm.

Holotype male: Carrville, Trinity County, California, 2,400 to
2,500 feet altitude. May 22, 1934 (G. E. Bohart). One male para-^
type, same locality and collector, June 18, 1934.

The holotype is in the California Academy of Sciences, San Fran-
cisco, California.

Proteriades deserticola new species

This is a robust, medium-sized species with abundant pale pubes-
cence. It is at least superficially similar to bidenticauda in having
two teeth on the seventh metasomal tergum of the male. It seems
probable, in view of the differences between both males and females,
that this condition arose independently in the two species and that
the bidentate male abdomen does not indicate close relationship.
In deserticola the two teeth are separated by a broad crescentiform
emargination and in bidenticauda by an equally broad quadrate

Bees of the Genus Proteriades 431

emargination. The female of deserticola has a very distinctive cly-
peal margin somewhat similar to that of evansi, but with the median
lobe less arched and more projecting over base of labrum, so that
the margin is about equally produced at sides and in the middle.

Male: Black, with first three metasomal terga usually entirely
red, except spot at base of first tergum (and more rarely a median
spot at base of second and third). Fourth tergum red except broad
median black patch; fifth and sixth terga black, red subapically and
broadly translucent ferruginous apically. Seventh tergum red.
Venter dark, variably suffused with red. Flagellum reddened be-
neath and tarsi more or less ferruginous apically. Clypeus with a
conspicous emargination as wide as labrum and limited by strong
angles, a small projection in middle of emargination. Mandibles
dark red on apical half, the teeth (sometimes) piceous. Tegulae
clear amber color. Wings clear hyaline, the nervures and stigma
brown. Head as broad as thorax, the genal areas about half as
wide as eyes seen from side. Inner orbits of eyes parallel. Posterior
ocelli slightly closer to eye margins than to the occipital margin.
Mandibles bidentate, the teeth subequal. Antennae slender, the
middle segments of flagellum barely longer than thick. Maxillary
palpi four-segmented. Middle coxae not toothed, posterior not
lamellate. Sixth metasomal tergum slightly toothed on each side;
seventh with a short blunt tooth at apical comers, these teeth sepa-
rated by a broad crescentiform emargination, the apex being broader
than usual. (Broad also in hidenticauda, truicauda and xerophila.)
Second sternum with a broad transverse subapical swelling. Third
to fifth sterna each with a slight transverse swelling close to apical
margin, the margin and swelling both indented medially. Sixth
sternum with high carina-like median fold. Puncturation very fine
and close on head and thorax, the punctures of vertex slightly closer
than those of mesoscutum. Abdomen slightly dullish, with minute,
close, shallow punctures. Pubescence white, dense on head and
thorax, with vertex, middle of mesoscutum and middle of propodeum
exposed. Vertex and exposed part of mesoscutum with short fine
hair (tinged with ochreous brown in fresh specimens). First four
metasomal terga each with a broad white apical hair-band, a little
narrowed in middle on first two. Fifth and sixth terga nearly uni-
formly covered with appressed white hair. Second to fifth sterna
each with a long white apical fringe, those on third to fifth conform-
ing to the curvature of the margin. Length, 5.0 to 6.0 mm.; fore
wing, 3.8 to 3.9 mm.

432 The University Science Bulletin

Female: Similar to the male. Flagellum beneath, except at base,
mandibles apically, and three basal metasomal terga ferruginous
red, the red usually a little darker than in male. Median sulcus on
first tergum and transverse streaks across bases of second and third
terga (usually) black. Head as broad as thorax, genal areas nearly
equaling the eyes in width as seen from side. Clypeal margin pro-
duced nearly equally at sides and in the middle, with the broad shal-
low emargination on each side of middle slightly bisinuate to form
a small vaguely indicated tooth on each side of the bluntly angular
median lobe, the latter rather well elevated, slightly projecting over
middle of base of labrum and arched as seen from in front. Mandi-
bles very stout, with inner margin parallel to outer, except for a
small emargination in inner margin near base; the two outer apical
teeth acute, inner tooth much shorter and only slightly acute, form-
ing nearly a right angle with inner margin of mandible. Hyposto-
mal carinae rather high behind and rather abruptly reduced at
angles, oblique anterior part of each low and but little shorter than
longitudinal part. Ocelli a little smaller than in male, the posterior
pair a little closer to edge of vertex than to the eyes. Puncturation
fine and close on head and thorax, and minute on abdomen. Pubes-
cence dense on face, nearly to level of ocelli, on cheeks, sides of
thorax and around margins of mesonotum, and white, except tinged
with brown, or brownish ochreous, on vertex and mesonotum. Hy-
postomal areas and mandibles with fringes of long hairs curving
over proboscidial fossa; long hairs also projecting from margin of
clypeus and from emargination of inner margin of mandible. Hair
of mesoscutum short, appressed, with slightly longer hairs inter-
spaced. Hair of scutellum, especially at sides and apex, and that
of metanotum much longer than that elsewhere on thorax. First
five metasomal terga each with a broad, dense, white apical hair-
band ; disks of fourth to sixth terga covered with not much thinner
appressed whitish hair. Legs densely covered with white hair on
outer and posterior surfaces of femora and tibiae. Ventral scopa
short and ochreous white. Length, 5.0 to 6.0 mm.; fore wing, 3.5 to
4.0 mm.

Holotype male: Palm Springs, Riverside County, California, on
Cryptantha barbigera, March 21, 1932 (P. H. Timberlake). Allo-
type female from the same locality and flower, March 30, 1945 (P.
H. Timberlake). Thirty-four female and eight male paratypes
from the same locality and flower, March 21 and 26, 1932, March
24, 1933, and March 30, 1945; nine female and one male paratypes

Bees of the Genus Proteriades 433

from the same locality, April 16, 1939 (R. ]\1. Boliart I ; one male
paratype, Indio, California, March 25, 1937 (R. M. and G. E. Bo-
hart) ; two female and six male paratypes, Andreas Canyon, Palm
Springs, Riverside County, California, April 7, 1940 (M. R. and R.
M. Bohart) ; t.vo male and one female paratypes, San Felipe Creek,
Imperial County, California, April 8, 1939 (R. M. Bohart); one
female paratype, four miles east of Edom, California, April 17.
1937 (T. D. A._Cockerell) ; two female paratypes, Thousand Palms
Canyon, California, on Cryptantha angustifolia, April 10, 1937 (P.
H. Timberlake, G. C. Varley); one male and one female, same lo-
cality, March 9, 1940 (R. M. Bohart); and one female paratype,
nineteen miles west of Kane Springs, California, on Cryptantha an-
gustifolia, March 25, 1933 (P. H. Timberlake); one male and one
female paratype, Borego, San Diego County, California, April 8,
1939 (R. M. Bohart).

The holotype and allotype are in the collection of the Citrus
Experiment Station, Riverside, California.

Proteriades evansi Michener

Proteriades evaiisi Michener. 1936, Bull, .'^outliern Cal. Acad. Sci., 35 : 92.

This species is most closely related to deserticola from which it
differs in the less extensive red areas, the acute inner mandibular
tooth, and the unproduced median apical portion of the clypeus
which does not overhang the base of the labrum, among other char-
acters. This species is known only in the female.

Female: Black, large, dorso-lateral areas on first metasomal
tergum, sometimes meeting dorsally, smaller lateral areas on second
tergum, and sometimes still smaller ones on third red. Posterior
margins of terga testaceous. Mandibles largely reddish, most
strongly so subapically. Antennal fiagellum dark brown above,
light brown beneath. Legs brownish black, distal segments of tarsi
paler. Tegulae testaceous. Wings dusky, veins and stigma brown-
ish black. Head slightly wider than thorax. Genal areas distinctly
wider than eyes seen from side. Distance between posterior ocelli
markedly less than distance from one of them to posterior margin
of vertex or to eye margin. Inner orbits slightly diverging below.
Clypeus low and broad, about three times as broad as long, not
bulging, depressed lateral lobes extending farther downward than
any other point, median part of lower margin much thickened and
slightly produced to form a very small truncation separated from
lateral lobes by broad shallow emarginations. Mandibles thick,
27—90

434 The University Science Bulletin

but little narrowed near bases, but with a small, well defined, and
deep subapical emargination of inner margin; apical teeth acute.
Hypostomal carinae high behind the obtuse angles, low in front of
them, the portions in front of angles about as long as those behind.
Maxillary palpi three- or four-segmented. Punctation fine, slightly
coarser in central part of clypeus than elsewhere on head, slightly
coarser on mesepisterna than elsewhere on thorax. Pubescence
sparse, forming weak apical bands at sides of first four terga, scat-
tered on fifth and sixth terga. Hypostomal areas with long hairs
curved toward mouthparts; clypeus with several long yellowish
hairs arising beneath margin of median truncation. Length, 6.6 to
7.6 mm.; fore wing, 4.2 to 4.6 mm.

The type of this species from Loyds, Tulare County, California
(August 3), is on deposit at the California Academy of Sciences.
Additional specimens are from Coffee Creek, Tulare County, Cali-
fornia, on Cryptantha jiaccida, June 11, 1925 ( P. H. Timberlake).

Proteriades seminibra (Cockerell)

Heriades scniinibm Cockeiell, 1898, Trans. Amer. Ent. Soc, 25: 198; Cockerell, 1903,
Psyche, 10: 76; Cresson, 1928, Mem. Amer. Ent. Soc, 5: 71.

Proteriades seminibra, Titii.'i, 1904, Jour. New York Ent. Soc, 12: 25.
Eriades seriuruber, Friese, 1911, Das Tierreich, 28: 29.

This species may be immediately recognized in the female by the
two long fingerlike processes projecting downward from the clypeal
margin. The male is far less distinctive, but can be recognized by
the small median prominence of the clypeal margin (a feature shared
with deserticola, which has the apex of the male abdomen bidentate).

Male: Black, first three metasomal terga red, the basal concavity
of the first infuscated, the third frequently and the second occasion-
ally infuscated middorsally; fourth and sometimes fifth terga red
laterally; sterna largely red. Mandibles red subapically. Flagel-
lum and tarsi brownish. Tegulae amber colored. Wings dusky
hyaline, the veins and stigma dark brown. Head broader than
long, as broad as thorax, genal areas about two thirds as broad as
eyes seen from side. Distance between posterior ocelli slightly less
than distance from one of them to eye margin or to posterior edge
of vertex. Inner orbits of eyes subparallel. Antennae long, all
flagellar segments longer than broad. Clypeal truncation broader
than width of labrum, demarked by distinct angles, gently concave
with margin impunctate and thickened, produced to an acute angle
just mesad of each end of truncation and to a blunt median angle.
Mandibular teeth acute, outer one somewhat longer tiian iimer.

Bees of the Genus Proteriades 435

Maxillary palpi rather elongate, four segmented. Middle coxae not
toothed; posterior coxae not lamellate. Sixth metasomal tergum
with a small but distinct angle on each side, seventh with lateral
margins not emarginate, each produced to ventral subapical angle,
posterior margin with broad emargination. First metastomal sternum
tiiickened subapically and with dense white pubescence on the poste-
rior declivity of this thickening. Second sternum with strong contin-
uous transverse convexity. Second to fifth sterna with long apical
fringes, those of third to fifth conforming to the broad emargina-
tions of the sternal margins. Sixth sternum with the usual fer-
ruginous median fold, the summit of which is flattened nearly to
the apex and bears short hairs. Head and thorax very finely punc-
tured, the scutellum and mesepisterna conspicuously more coarsely
so than head and mesoscutum. Abdomen minutely and closely
punctured. Pubescence white, that of face above clypeus, vertex,
and dorsum of thorax ochraceous, that of clypeus exceedingly dense.
Pubescence of abdomen short, forming apical white bands at sides
of first four metasomal terga, but these bands apparently easily
worn off. Length, 5.8 to 7.5 mm. ; fore wing, 4.2 to 4.8 mm.

Female: Similar to male in color and sculpture. Black, first three
metasomal terga red (darker than in male), except a medial basal
patch on each segment, that on the third usually very large. Mandi-
bles variably red at apices and on inner margins. Flagellum strongly
reddened beneath, and small joints of tarsi sometimes ferruginous.
Tegulae amber color. Wings dusky, with blackish stigma and
nervures. Head large and massive, usually much wider than thorax.
Genal areas broader than width of eyes and rounded behind. Dis-
tance between posterior ocelli much less than distance from one of
them to nearest eye margin or to posterior margin of vertex. Clyp-
eus convex, much produced over base of labrum. Anterior margin
of clypeus with a small rounded lobe far to each side and in the
middle a large projecting lobe provided with a fingerlike process
on each side, between which the margin comes to an acute point,
the fingerlike processes being strongly divergent. Mandibles large,
with the apical two thirds of each cjuadrate, with nearly parallel
inner and outer margins, this portion being about one and one fourth
times longer than wide. Inner margin of mandible deeply excavated
near base, as is also the exterior surface as far outward as the inner
mandibular carina, this excavation making room for the rounded
lateral lobes of the clypeal margin, when the mandibles are closed.
Mandibular teeth equal, acute. Antennae much shorter than in

436 The University Science Bulletin

male, the flagellum cylindrical, thinner than the scape and about
twice as long. Hypostomal carina well elevated, behind the angle,
low in front of it, the part anterior to the obtuse angle very obliciue.
the two parts being nearly equal in length. Almost entire surface
with fine, very close punctures, but, as in male, the large basal area
of propodeum is smooth and sculptureless except for a finely rough-
ened strip across the base. Pubescence whitish, moderately dense
on face, nearly to level of ocelli, and often tinged with brown in
fresh specimens on vertex and notum of thorax. First five meta-
somal terga each with a dense white apical hair-band, more or less
thin or interrupted in the middle on first two terga. Disks of last
two segments rather densely covered with appressed whitish hair,
and disk of preceding segments with short erect hair. Length, 5.8
to 8.5 mm. ; fore wing, 3.8 to 5.0 mm.

Cockerell's two type specimens of scDiirubra were from southern
California, and Titus had a male from Los Angeles County collected
by Coquillett. This probably is the commonest species of the genus
in southern California, although at higher elevations it is replaced
in this respect by jacintana and on the deserts by xerophila and
deserticola. At Riverside the senior author has taken males from
March 22 to May 18 and females from April 4 to .June 7. It fre-
quents spots only where Cryptantha intermedia is growing, and the
female obtains its pollen exclusively from the fiowers of this plant.

Additional locality records (all in California) are Ferris; the
Gavilan, Riverside County; six miles east of Temecula, Riverside
County; Warner's Hot Springs, San Diego County; Verdemont, San
Bernardino County; Mill Creek, San Bernardino Mountains (all
collected by P. H. Timberlake). Redlands, San Bernardino County;
Andreas Canyon, near Falm Springs; San Jacinto River, San .Ja-
cinto Mountains, 4,000 feet altitude; Newhall, Los Angeles County;
Coalinga ; .Jamesburg, Santa I^ucia Mountains, Monterey County
(all collected by C. D. Michener). Pinacles National Monument,
San Benito County (F. D. Hurd). Atascadero, San Luis Obispo
County (E. C. Van Dyke). All specimens were collected within the
range of dates indicated for Riverside, except that in Monterey
County males were collected as late as June 4 and females as late
as June 8. All specimens for which comjilete flower records exist
were on Cryptantha intermeuia; some labeled merely "Cryptantha"
may have been taken on other species of the genus. A total of over
200 specimens of this species have been examined.

The lectotype male is in the collection of the Academy of Natural

^ Bees of the Genus Proteriades 437

Sciences of Philadelphia. A neallotype female, here designated, is
from Redlands, California, May 15, 1936 (C. D. Michener), in the
Snow Entomological Collections.

Proteriades jacintana (Cockerell) new combination

Chelostoma fCephalapis) jacmtaiia Cockerell, 1910, Ann. Mag. Nat. Hist., (8) 5: 23.
Cephalapis jacintmm Cockerell. 1925. Proc. California Acad. Sci.. (4) 14: 200.

This species is very easily recognized by its elongate form, broad
genal areas, the keeled clypeus, the large and virtually tridentate
male mandibles, and the long and nearly straight margin between
the upper two teeth of the female mandibles.

Male: Black, the first two to four (usually three) metasomal
terga red at sides, the red of the first tergum most extensive and
often forming a band across the posterior margin of this tergum;
first six terga with posterior margins brownish. Mandibles rufescent
subapically. Under surface of antennal flagellum often dark brown.
Tegulae amber colored. Wings slightly dusky, veins and stigma
brownish black. Genal areas as wide as eyes seen from side. Dis-
tance between posterior ocelli slightly less than or subequal to dis-
tance from one of them to eye margin, much less than distance to
posterior margin of vertex. Inner orbits of eyes parallel above, di-
verging below. Antennal flagellum with middle segments longer
than broad. Mandibles large, curved, apices tridentate, outer tooth
much longer than next, outer two both acute, inner tooth widely
separated from outer two, right angular. Clypeus thickened, with
a longitudinal median keel the lower end of which ends at a slight
median projection of the produced truncation; truncation thickened,
much overhanging base of labrum, short and demarked laterally by
angles of about 90°. Maxillary palpi four-segmented. Hypostomal
carinae strongly elevated posteriorly. Middle coxae each with
a distinct ventral tooth in front of base of trochanter. Posterior
coxae each with carina elevated so that it might be considered a
low lamella. Sixth abdominal tergum with a rounded angle at each
side. Seventh tergum deeply emarginate at apex, the resulting lobes
longer than wide, blunt, each about as broad as space between them.
First metasomal sternum somewhat thickened with declivous, hairy,
posterior margin. Second sternum large, weakly fringed. Third to
fifth sterna well fringed, fringe of third dense in middle, of fifth con-
forming to broadly emarginate sternal margin. Sixth sternum with
the usual ferruginous fold very long and well developed, ending
rather abruptly at base, terminating in a short spine, summit of fold
broadened and provided with short hairs. . Punctation fine and

438 The University Science Bulletin

dense, coarser on mesepisterna than elsewhere on head and thorax.
Pubescence rather dense, white, slightly ochraceous on dorsum of
head and thorax, dense on face nearly to level of anterior ocellus,
forming apical bands on first five metasomal terga. Length, 6.5 to
8.4 mm. ; fore wing, 4.5 to 5.2 mm.

Female: Coloration similar to male, but red confined to first two
metasomal terga and not extending as a band across posterior mar-
gin of first. Genal areas much broader than eyes seen from side.
Distance between posterior ocelli less than distance from one of
them to eye margin, much less than distance from one of them to
posterior margin of vertex. Inner orbits diverging below. Clypeus
W'ith a carinate longitudinal median keel; truncation much over-
hanging base of labrum, much shorter than distance from end of
truncation to lateral angle of clypeus; sublateral angle w^eak and
midway between end of truncation and lateral angle of clypeus.
Mandibles large, narrowed near bases by emarginations of upper and
lower margins, mucli expanded apically, middle and inner teeth
scarcely acute, wadely separated, connected by long and nearly
straight margin. Maxillary palpi four-segmented. Hypostomal
carinae elevated posteriorly. Carinae of posterior coxae reduced
and inconspicuous. Punctation fine and dense, coarser on clypeus
than on rest of head. Pubescence sparser on face than in male, form-
ing apical bands on first four terga, scattered, denser apically, on
fifth and largely covering sixth. Length, 7.0 to 10.5 mm.; fore wing
4.2 to 5.4 mm.

The male type of this species, from Kenworthy, San Jacinto Moun-
tains, California, is in the collection of the Citrus Experiment Sta-
tion at Riverside, California. The species is a common one, es-
pecially in the southern California mountains and is know-n from
the following localities, all in California: Warner's Hot Springs,
San Diego County; Stone Creek, Keen Camp, Idyllwild, San Jacinto
River (4,000 feet altitude) two miles southeast of Poppet Flat.
Herkey Creek, Vandevanter Flat, Ribbonwood, and Mount Santa
Rose (7,500 feet altitude), all in San Jacinto Mountains; Mill
Creek (4,400 to 4,700 feet altitude) and Forest Home, both in
San Bernardino Mountains; Ortega Highway, Santa Ana Moun-
tains, overlooking Elsinore; Riverside; Claremont; Eagle Rock,
Los Angeles County; Newhall, Los Angeles County; Pinacles, San
Benito County; Jamesburg, Santa Lucia Mountains, Monterey
County (1,900-2,700 feet altitude); Yosemite (3,880-4,000 feet al-
titude); Midway; Madera County (3,000 feet altitude); Coffee

Bees of the Genus Proteriades 439

Creek and Carrville (2,400-2,500 feet altitude), Trinity County;

Big Pine Creek, Inyo County (7,500 feet altitude) ; Rock Creek,

Mono County (8,500 feet altitude). The flower visited in southern

California is usually Cryptantha intermedia but it has also been

collected on C. micrantha var. lepida. Flower records for the more

northern collections are merely Cryptantha sp.? Dates of collection

for males range from April 20 to June 23, for females from April 20

to July 5.

LITERATURE CITED
MicHENER, Charles D.

1941 A generic revision of American Osmiinae with descriptive notes on
Old World genera (Hvmenoptera, Megachilidae), Anier. Midland
Nat., 26: 147-167.

1943 American bees of the genus Anthocopa with notes on Old World
subgenera (Hymenoptera, Megachilidae), Ann. Ent. Soc. Amer.,
36: 49-86.

1944 Comparative external morphology, phvlogeny, and a clas.*;ification
of the bees (Hymenoptera), Bull. Amer." Mus. Nat. Hist.. 82: 151-326.

1944a The distribution of the osmiine bees of the deserts of North America.
Amer. Nat., 78: 257-266.

44U

The University Science Bulletin

PLATE III

.i^r'^^^^^ii^

1-6, Antennae of males, 1. /'. foninilna ; 2. P. boliarti ; 3, P. caudex ; 1. P.
bidenticauda ; 5, P. hamuliconiis; 6, /-". pygmacn. 7-9, Apical portions of
mandibles. 7, P. jacintana, female; 8, P. jncintana, male; 9, P. semirubm,
female. 10-16, Sixth and seventh metasomal terga of male. 10, P. jariutnnn;
11, P. bidenticauda; 12, P. scDiirnbrii ; 13, /•'. Ii(nuulicnntis ; 14. P. bnhart? ;
1."), P. r-ii(l(.r; 16, P. pyfjuiacd.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. 11] March 20, 1950 [No. 11

Second Contribution to the Herpetology of
San Luis Potosi

BY

Edward H. Taylor,

Department of Zoology, University of Kansas

Abstract: A report i.s made on a small herpetological collection from the
Mexican State, San Luis Potosi. One new species, Adelphicos neionuutorum,
is described, and the known ranges of several ^pecies are extended to th'"i
State.

In a recent issue of this journal,* thanks to the kindness of Prof.
George H. Lowery of Louisiana State University, I reported on a
hirge herpetological collection made in the state of San Luis Potosi,
Mexico, by a collecting party from the Louisiana State University,
consisting of Mr. and Mrs. Robert Newman and Mr. Charles Shaw.
The bulk of the collection, however, was acquired by Mrs. Newman.
Most of the specimens were taken in the region of Xilitla, in the
southeastern portion of the state; a few, however, were obtained in
other localities.

One group of specimens from the collection was described by Dr.
J. A. Tihen as a new subspecies, Gerrhonotus leiocephalus loweryi.'f

On July 2, 1948, a second collection was forwarded to me from
Louisiana State LTniversity. This material was likewise acquired in
San Luis Potosi, largely in the Xilitla region, by a collecting party
consisting of Mr. and Mrs. Robert Newman, Mr. Herbert Stern, Mr.
Prentiss D. Lewis, and Mr. George Lowery. The material, while
not so extensive as the preceding collection, does contain material
of importance, including certain new state records. One of these
is a very rare Mexican rattlesnake, Crotohis basiliscus totonacus.

* Univ. Kansas Sci. Bull., vol. 33, pt. 1, pp. 169-215.

t Trans. Kansas Acad. Sci., vol. '>\, 1948, r.p. 302-30;").

(441)

442 The University Science Bulletin

Altogether there are 55 specimens representing 23 species. I am
again obligated to Professor Lowery for the privilege of examining
this second collection.

The following species, one of which is described as new, are here
recorded from San Luis Potosi for the first time:
Adelphicos newmanorum sp. nov.
Thamnophis subcarinatus subcarinatus (Gray)
Drymobius chloroticus (Cope)
Micrurus fulvius tenere (Baird and Girard)
Crotalus basilisciis totonacus Gloyd and Kauffeld.
The entire known herpetological fauna of San Luis Potosi totals
140 species. This is divided as follows: salamanders, 5; Salientia,
25; lizards, 33; snakes, 78; turtles, 3.

The map showing collecting localities in San Lui'S Potosi, as well
as the data on the localities included in the text, has been prepared
by Dr. George H. Lowery to whom I offer my sincere thanks.

Amphibians

Bufo valliceps Wiegmann

A single specimen having the typical light coloration is in the col-
lection.

Smilisca baudin'i Dumcril and Bibron

A single medium-sized female specimen was taken.

Lizards

Sceloporus variabilis ; ar/a 5//;'.s Wiegmann

A series of eight specimens was taken at Hda. Limon, 10 mi. W of
fibano, San Luis Potosi between March 30 and April 24, 1948.

Sceloporus serrifer plioporus Smith

A single specimen was taken at Hda. Limon, 10 mi. \V. of fibano.
The femoral pores are 20; the dorsal scales, 31. The color, and
much of the pattern, has disappeared due to jn-eservation.

Sceloporus granimiciis disparilis Stejneger

A single specimen taken on the Llano de Coneja at 7,000 ft., is
referred to this subspecies. The dorsal scales number 70 from occi-
put to a line joining the posterior surface of the thighs. This
locality is near the area of intergradation between g. disparilis and
g. microlepidotus.

Taylor: Herpetology of San Luis Potosi 443

Corythophanes hernandesii (Gray)

Seven specimens, Nos. 615 to 621, are in the collection, all from
the Xilitla region. These were obtained by Marcella Newman.

Gerrhonotus leiocephalus loweryi Tihen

Three topotypic specimens from Xilitla are in the collection. Two
(Nos. 612 and 614) have the granular area in the lateral fold defi-
nitely marked with vertical dark lines. The smallest specimen (No.
613) shows no such markings. Otherwise they agree in significant
characters.

Ameiva undulata podarga Smith

Five specimens (Nos. 623-627) are in the collection, four from
Xilitla and one (No. 627) from Tamuzunchale.

Snakes

Ninia dindcmata plorator Smith

A portion of a badly jn-eserved specimen from Xilitla is included
in the collection (No. 609).

Adelphicos neivmanorum, sp. nov.

PI. IV

Holotype: Louisiana State University No. 204, ^ , Xilitla region,
San Luis Potosi, May 7, 1947; Marcella Newman, collector.

Diagnosis: A species of average size for the genus having the
third labial replaced on edge of lower lip by the enlarged first chin-
shields; no lateral stripes present on the body; venter immaculate
white; subcaudal region with a slight median peppering of pigment;
above brownish gray, the scales faintly outlined with heavier pig-
ment.

Description of type: Rostral at least one third wider than high,
the part visible above triangular, as long as the suture between the
internasals; latter scales small, wider than long, their length about
one third the length of prefrontals, their area between one fourth
and one fifth of the prefrontals; prefrontals as broad as long, en-
tering the orbit; frontal triangular, as long as wide, about one fifth
longer than the prefrontal; parietals elongate, about one fifth longer
than their distance from the end of the snout, their common suture,
however, being about two thirds the length of the frontal; nasal
divided, the anterior part less than half the area of the posterior,
the nostril pierced between the two parts very close to the rostral;
loreal slender, elongate, entering orbit on right side, while on left

m

w-

I'late IV. Fig. 1. Adelphicos newmanorum sp. nov. Louisiana Stato
University No. 204, Type. Xilitla region. San Luis Potosi, Mexico
(U)tal length, 310 mm.). Fig. 2. Adclphicos quadrivirgatu-'i quadrivir
{/(it us Jan. Louisiana State University No. 206, Xilitla, San Lui^
Potosi. Mexico (total length, approx. 360 mm.).

Taylor: Hp:rpetology of San Luis Potosi 445

side, shorter, triangular, excluded from eye by contact of third
labial and prefrontal; no preocular; two postoculars, the upper
double size of lower; temporals large, one behind the other, the
posterior as large or slightly larger than anterior.

Supralabials, 7-6, in the following ascending order of size: left —
1, 2, 3, 6, 4, 5, 7, the third and fourth entering the orbit; on right
side the second and third are fused, and only the fourth enters the
the orbit; infralabials presumably 6, the third being replaced by the
much widened first chinshield, first pair of chinshields followed by
an azygous pregular; second pair of chinshields not or scarcely dif-
ferentiated, and separated by a second median gular a little larger
than the first. Diameter of eye about 2V1> times in length of snout.

Scale formula: 15-15-15; ventrals, 139; caudals, 49; anal divided;
total length, 310 mm.; tail 61 mm.; head length, 7.5 mm.; head
width, 5.9 mm.

Color in jormalin: Above light grayisli brown, the edges of all
scales somewhat more heavily pigmented than centers; a very faint
median grayish line; venter and outer scale row white; second scale
row light but with some scattered pigment; an indistinct gray sub-
caudal line; head dark reddish brown; labials, save for their upper
edges, cream.

Relationship : This form is related to quadrivirgatus quadrivirga-
tus (PI. IV, fig. 2) in having the anterior chinshield form a part of
the labial border and in having the chinshield followed by two
median gulars, one following the other, separating the poorly differ-
entiated second pair of chinshields.

The head, however, is somewhat more slender than in that species,
the frontal being longer than its distance from the tip of the snout
(in quadrivirgatus less than this same distance). The top of the
head is more reddish brown, and the labials are somewhat lower.
The typical quacrilinear markings of q. quadrivittatus are lacking.

Remarks: On one side the lower jaw, and an area on the neck
has been injured by ants.

Bocourt* has described a slender-headed variety of Adelphicos
quadrivirgatum as acutirostrum (type locality, "Mexique"). His
form was striped. The subcaudal count was 32, the ventrals, 135.
Smith and Taylor (Bull. 187, U. S. Nat. Mus., p. 30) regard this
form as a synonym of Adelphicos q. quadrivirgatus Jan.

* Mision Scientifique au Mexique . . ., fitudes sur les Reptiles, livr. 9 1883 pi 32
figs, lib, lie, 12b, 12c.

Plate V. Fig. 1. Ficimia streckeri Taylor; Louisiana State University No.
214 S , Xilitla, San Luis Potosi, Mexico (total length, 426 mm.). Fig. 2. Ficimia
streckeri; Louisiana State University No 208 $ , Xilitla, San Luis Potosi, Mexico
(total length, 383 mm.).

Taylor: Herpetology of San Luis Potosi

447

Plate VI. Fig. 1. Rhadinaea forbesi Smith; paratype. Fig. 2. Rhadinaea
mnrcellac Taylor. Louisiana State University No. 270. Type, Xilitla region,
San Luis Potosi, Mexico (total length, 290 mm.).

448 The University Science Bulletin

Ficimia streckeri Taylor

PI. V. figs. 1 and 2

Three specimens in the collection from Xilitla, Nos. 605 ^ , 606 ^ ,
and 607 5 , respectively, yield the following scale data: postoculars,
2-1, 2-2, 1-1; scale formulae, 19-17-17, 19-17-17, 19-17-17 (the
number 19 occurs only on the anterior part of the neck) ; ventrals
156, 153, 142; caudals, 34, 34, 33; totals, 190, 186, 175. Most of
the scales on the anterior part of the body in the two younger speci-
mens have single median terminal pits instead of the typical pair.
In the older specimen, No. 607, however, the pits are scarcely dis-
cernible. If the epidermis has been shed they cannot be seen.

The narrow bars, present in Nos. 606 and 607, to the number of
41 and 43 respectively, are, in No. 605, replaced by a median series
of spots, two or two and one half scales long and about as wide.
There are but thirty such marks. On the sides are two rows of
elongate spots below which are some smaller indefinite spots reach-
ing the ventrals.

Rhadinaea marcellae Taylor
PI. ^•l, fig. 2

Rhadinaea marcellae Taylur, Univ. Kan.sas. Sri. Bull., vol. 3.S, pt. 1, ]949, pp. 197-198.

The type specimen of this species is figured. A figure of Rha-
dinaea forbesi Smith is also included for comparison. (PI. VI, fig. 1.)

Rhadinaea crassa Smith

Three specimens, Nos. 601-603, of this species, two from Xilitla,
and one from Cueva Salitre, near Xilitla, are in the collection. The
tails of Nos. 601 and 603 are broken, while the body of No. 602 is
badly crushed.

Elaphe flavirufa fiavirufa (Cope)

A very large male specimen, No. 585, was collected at Hda.
Limon, 10 mi. W of Ebano, San Luis Potosi, by Prentiss D. Lewis.
The specimen measures 1525 mm. in total length, the tail 286 mm.

This specimen dilTers from younger specimens in having scattered
ventral black flecks (of a size equal to the surface area of one or
two dorsal scales) that grow more numerous on the jiosterior part
of the body, so that the dark color rather than the light is pre-
dominant. The subcaudals are dark over most of their surface, only
a small anterior portion t)f each being white.

There are 34 dorsal body spots and 15 caudal spots; the ventrals
are 247; anal divided; caudals, 106.

Tayl(jh: Hi:RPKTOLO(iY OF San Luis Potosi

449

M0^

■-»

,♦

Plate VII. Pliocercus laticollaris Smith, Louisiana State University No. 559.
Xilitla, San Luis Potosi, Mexifo (total length, 541 mm.).
28—90

450 The University Science Bulletin

Pliocercus laticollaris Smith

PI. VII

A specimen of this species was reported from Xilitla in my first
paper on the San Luis Potosi fauna. A figure based on a photo-
grajih of tlie jireserved specimen is inchided here.

Dnjmobiiis chlorot'cus (Cope)

Dendrophidium chloroticum Cope, Proc. Amer. Phil. Soc, vol. 23, 1886, p. 278.
Drgmohiun chloroticus Cope, U. S. Nat. Mus. Bull., no 32, 1887, p. 09.

This species, with a type locality at Coban, Guatemala, has here-
tofore been reported in Mexico only in southern Veracruz and in
Chiapas. The collection contains one specimen, No. 591, young 5 ,
collected at Xilitla, San Luis Potosi, by Mr. Herbert Stern, Jr.
This is the most northern record, extending the known range about
280 miles to the northwest.

The ventrals are 164; anal divided; subcaudals, 116 -\- 1.

The original green coloration has disappeared and a dim pattern
of transverse darker marks, narrowly edged with white, is evident;
the whitish edges of the bands are separated by three scales. An-
teriorly, the skin between the scales is light. The head is black
but most of the rostral and the supralabials are cream except on
their upper edges. The infralabials and chin are cream white. The
coloration of the sides extends onto the ventrals. There are 38
maxillary teeth, the posterior approximately a half larger than
anterior. The scale formula is 17-17-15, the eleven middle scale
rows with keels.

Drymobius margaritiferus margaritifcrus (Schlegel)

A single specimen, No. 587, was collected at Hda. Limon, ten mi.
W of Ebano.

Dryadophis 7nektnolo}nus veraecrucis Stuart

Six specimens, Nos. 593-598, are in the collection; two are from
Apetsco, four from Xilitla. A young specimen, No. 598, has the
body uniform dark brown. A white line beginning on the labials
below the eye can be traced along the side of the neck and body
where it appears so dim that it can scarcely be traced. I cannot dis-
cern any transverse markings in this specimen.

Spilotes puUati^s me.vicnni)fi (Laurcnti)

A young specimen. No. 586, is in the collection from Xilitla. The
head has the reduced pattern of black. The dark bars on the body,
after the first eleven, are nearly transverse losing the strong diagonal

Taylor: Herpetology of Sax Luis Poxosi 451

trend of the anterior bars. There are 21 bars on the body posterior
to the nuchal band, and 12 caudal bands. Ventrals, 200 ; anal
single; caudals 118 -f 1.

Larnpropeltis triangidum polyzona Cope

Two female specimens, one from Apetsco, Xilitla Region (No.
588), and one from Xilitla (No. 589), are referred to this subspecies.
One of the specimens has the black bands widened dorsally so that
in many cases the bands are separated on the middorsal line by a
single row of red scales. This specimen might be regarded as an
intergrade with the subspecies arcifera known also from the Xilitla
region. In this specimen the red scales are shaded with dark pig-
ment but there are no distinct black spots.

No. 588 from Apetsco is a typical polyzona, the red interspaces
being as wide as the black bars. Each red scale has a discrete
black spot.

The ventral and subcaudal counts for Nos. 588 and 589 respec-
tively are: 211, 216, 46 + 1, 49 + 1.

Leptodeira annulata septentrionalis (Kennicott)

This species is represented by a single specimen (No. 592) from
Xilitla, taken April 25 by Marcella Newman.

The specimen yields the following scale data: Preoculars, 3-3, the
upper touching the frontal on each side; postoculars, 2-2; supra-
labials, 8-8; infralabials, 10-10; temporals, 1 + 2; scale formula
21-23-23-17; ventrals, 195; caudals, 73; anal divided. Thirty bands
on body; 16+ on tail.

The minute grayish pigment flecks on the venter are fewer than
those usually present. The bands are a little narrower than those on
specimens from farther north.

Thamnophis subcarinatus subcarinatus (Gray)

Coluber {Xatrii) subcaririata Ciray, Zoologj' of Beechey's Voyage, 1839, p. 9(i, i)l. :i2 (t\iu'
locality "the hedges of Xalisco" Mexico.)

Thatniioiyhiii fnibrarinaia subcariiiuta Smith, Herpetologica, 1949, vol. 5, pp. (i3-<i4. (Tyi)e
locality restricted to CJuadalajara. .Jalisco.)

This species is represented by a male specimen (No. 5901 col-
lected at Laguna de las Rusias, Villa de Reyes region, San Luis
Potosi. The ventrals are 163, the caudals 64 -j- 1.

The reduced caudals in the male separates this subspecies from
subcarinatus megalops that has 79 or more subcaudals. The latter,
a more western subspecies, has also been found within the limits
of the state.

452

The University Science Bulletin

Storeria dekayi texana Trapido

A specimen from Xilitla (No. 608), with 17 scale vows in the mid-
dle of the body, two pairs of chinshields present, lacking both a
loreal and a horizontal dark mark on the anterior temporal, is re-
ferred to this species. The ventrals in this specimen are 139; anal
divided; and caudals 55. The scale formula is 17-17-17.

Plate VIII. Mirnuus fulvia.s tcnerc (Baiid aiul Girard.) Louif^iana State Uni-
versity No. 310. fibano, San Luis Potosi. Mexico (total length, 703 nun.).

Taylor: Herpetology of San Lns PoTosi 453

The dorsal coloration in preservative is nearly uniform brownish
gray with some few small blackish spots anteriorly. The venter is
lighter with two or more black dots on the end of each ventral. On
the underside of the tail the caudals are edged with gray and are
Avithout dots.

Micrurus fulvius tenere (Baird and Girard)

PI. VIII

This species is represented by a single specimen, L.S.U. No. 310.
I originally reluctantly referred* this specimen to Micrurus fitzingeri
microgalbineus Brown and Smith. I pointed out the characteristics
by which the specimen differed from typical M. /. microgalbineus,
and ga-\'e scale data. The colors are badly faded. A figure of the
specimen is included.

Micrurus fitzingeri microgalbineus Brown and Smith

Pi. IX

A figure of L.S.U. No. 308, Xilitla, San Luis Potosi, is given.
Crotalus basiliscus totonacus Gloyd and Kauffeld

Crotalus totonacus Gloj'd and Kauffeld, Bull. Chicago Acad. Sci., vol. 6, no. 2, p. 12, fig. 1-2
(type, Chicago Academy of Science no. 44&0, Panaco Island [about 75 mi. S of Tampico],
Veracruz, 12 mi. inland from Cabo Rojo).

Since only one specimen besides the type and paratype of this
species is known, and only the type has a definite locality, it is with
very considerable interest that I report the discovery of this rare
species on tiie mainland of Mexico at Hacienda Limon, 10 mi. W
of Ebano, San Luis Potosi. These specimens, Nos. 583 and 584, were
collected respectively by Mr. Herbert Stern, Jr., and Mr. Prentiss
D. Lewis.

The smaller complete specimen, No. 583, measures 813 mm. in
total length, the tail, 68 mm. The rattle is short, being only 16
mm. The head measurements are: width, 26.6 mm.; length. 33 mm.
The ventrals are 194, the caudals 26. The anal is single. The nos-
tril is higher than wide, and broadly in contact with the prenasal.
A pair of internasals are present, followed by a pair of prefrontals
(one prefrontal and one internasal being partially fused). The
supraoculars are large, flat, and separated anteriorly by a single
frontal that is divided transversely near the middle, and with the
posterior portion of the frontal apparently segmented longitudinally.
In contact with the posterior frontal and the supraoculars, is a some-

Univ. Kansas Sci. Bull. vol. 33, pt. 1, 1949, p. 213.

454

The University Science Bulletin

Plate IX. Micrunis fitzinr/cri micyogalblncns Brown and Smith, Loui«iana State
Tlnivensity No. 308, Xilitla, San Lui.s Poto^i, Mexico.

what enlarged parietal separated i'runi itt< fellow by a pair of scales.
The supralabials are 12-12, the infralabials 13-13. The nostril is
represented by a vertical slit between the nasals and the second
nasal is followed by a pair of superimposed loreals. Two large pre-
oculars follow these scales, both entering the orbit. A series of foin

Taylor: Herpetology of San Luis Potosi

455

diminutive scales are present anterior to the large scale bordering
the pit. The pit is surrounded by four scales, including the lower
preocular. The two lower scales are in contact with the labials.
There are two or three scale rows between the labials and the eye.
The three lower rows of temporals are unkeeled, the upper rows
slightly keeled. Three labials touch the anterior chinshields.

The color pattern of this specimen resembles closely the figure
of the type given by Gloyd (1940), even to the smaller detail of the
head pattern.

The second specimen is a large male, but the head, with a portion
of the neck, is missing. The preserved portion measures 1360 mm.
of which tail and rattle measure 150 mm., the rattle alone, 35 mm.
The scale formula is 25-25-23-21.

Bothrops atro.v asper (Garman)

Three specimens of this large species are in the collection (Nos.
599 to 601). Two large specimens are from Xilitla, a small one
from Hda. Limon, 10 mi. W of Ebano.

I

102 lOi 100 99

1 1

/ \

/

\

/

i SAN LUIS POTOSI

24

/

1

-

24

/

^

y

\

y

/

!

/

/

/~

'

/

v-.r--,

CHARCAS*

s

23

\

— % —

23

7

/

•"

■\

"n

\

s .-.

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PEN

3ENCIA» • \sABI|v»IT0

/CANDI

DO NAVARRO

• • — •-«._». -'■>^.

Uan L

• ^ Cll

JDAD DEL MAIZ . ^.

J IS POTOSI

•

EBANO w^.

22

] '

'EAJE

TAMU N • j

22

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•

i/ALLES Cj-

_»VILLA DE REYES

• pujAL ■^

0 10 25 50 100

1 1 1 1 1

\ — . _

CIUDAD !

5ANT0S^•
• \

III) 1

Kilometers

'^- i' ^

MuMum of Nolurol History

^'ct^Rd^.QP^JEJAV,

XILITLA^'

1 ,

Univortity of Konaoa
1947

TAMAZUNCHA

AJHUACAjTLAN

102 101 100 95

Map showing collecting localities in San Luis Potosi

456 The University Science Bulletin

The three specimens present the foHowing scale data:
Data on Bothrops alrox asper (Garman)

Xo.

Sex

Veiitriih

Suh-
caudah

/.

ahials

hijra-
kihials

Scale foriniild

Total
length

Tail
Iciipth

599

s

20.')

03 + 1

7-7

10-10

27-29-21 -20

1,535

184

COO

i

209

65 + 1

7-7

11-10

20-25-21-19

1,500

197

COl

5

211

01 + 1

7-7

10-10

28-2(1-21-20

1,015

132

LIST OF LOCALITIES

Ahuacatl.an: a small village at 3,800 feet situated among nunintains rising
to over 5,000 feet.

Apetsco: a locality api>roximateIy L5 miles NW of Xilitla; elevation rn.
2,600 feet.

AxTL.A Ferry on Ro.\d to Xilitla: a ferry crossing on that part of the Rio
Axtla locally known as the Rio Huichihuayan ; ele\ation en. 300 feet.

C.ANDiDO Nav.arro: a village on Ciudad San Luis Potosi-Antiguo Morelos High-
way approximately 15 miles northeast of the city of San Luis Potosi.

Cerro Coneja: the highest measured peak in the Sierra Madre Oriental
proper of San Luis Potosi; elevation 8,699 feet. Specimens with this
locality designation were taken at elevations ranging between 6,200 and
7,400 feet.

CiUDAD Del Maiz: a town on Ciudad San Luis Potosi-Antiguo Morelos High-
way; elevation 4,000 feet.

CiUDAD Santos: current name for village formerly known as Tancanhuitz;
elevation 770 feet.

Charcas: a town 68 miles north of Ciudad San Luis Potosi; elevation 6,580
feet.

Cueva Salitre: a cave located on the outskirts of village of Xilitla.

Ebano: a town near estern border of state; elevation ca. 100 feet.

El Salto: a waterfall about 5 miles N of Ciudad del Maiz-Antiguo Morelos
Highway; elevation 2,000 feet. Specimens were taken at an elevation of
about 1,500 feet. This is Salto del Agua of some maps.

Hacienda Limon: a general collecting site about 10 miles W of Ebano; elo-
vation 45 feet.

JoYA De La Silleta: a locality adjacent to Llano do Garzas; elevation at.
6,200 feet. The spelling of this place name was erroneously transliterated
in the early stages of the field work as "Joya de Asietta,"

L.va'NA De Las Ruslas: an artificially emjjounded body of water, with den.se
stands of rushes and other aquatics, located 3 miles SW of Villa de Reyes;
elevation 6,000 feet.

Llano De Garzas: one of several mountain meadows in neighborhood of Cerro
Coneja; elevation ca. 6,800 feet.

Llano De Coneja: a mountain meadow in neighborhood of Cerro Coneja;
elevation ca. 7,000 feet.

Miramar: this locality designation refers to Cerro Miramar, a mountain over-
looking the village of Xilitla; elevation ca. 4,500 feet.

Pea.ie: a village approximately 8 miles SW of the city of San Luis Potosi
on highway to Guadalajara; elo\ation ca. 6,400 feet.

Taylor: Herpetology of San Luis PoTOsi 457

Pexdexcia: a village 2.5 miles N of San Luis Potosi-Antiguo Moiolos High-
way; elevation ca. 4,500 feet.

PKE.SA De Guadalupe: a village on Ciudad San Luis Potosi-Antiguo Morelos
Highway; elevation ca. 4,000 feet.

Puerto De Lobos: a point on Ciudad San Luis Potosi-Antiguo Mon^los High-
way 2.5 miles S of Pendencia; elevation 4,300 feet.

Pujal: a town on Pan-American Highway; elevation ca. 300 feet.

Rancho Miramar Grande: a locality about halfway along trail between Cerro
Miramar and Cerro Coneja, including collecting points from 4,500 to 5,000
feet.

Sabinito: a village on Ciudad del Maiz-Antiguo Morelos Highway; elevation
L300 feet.

Sax Luis Poxosi Reservoir: an impounded body of water on the southwestern
outskirts of the city of San Luis Potosi; elevation 6.300 feet.

Tamazunchale : a town on Pan-American Highway in the southeastern part of
the State; elevation ca. 400 feet.

Tamuin: the present name for a village formerly known as Guerrero, located
on Valles-Tampico Highway; elevation 150 feet.

V.ALLEs: a town on Pan-American Highway; elevation 250 feet.

^'iLL.^ De Reyes: a town 25 miles S of the city of San Luis Potosi; elevation
6,000 feet.

Upper Llano De Coneja: a meadow on shoulder of Cerro Coneja; elevation
7.200 feet.

Xilitla: a town in southeastern part of State. 14 miles by road west of Pan-
. American Highway; elevation ca. 2,200 feet.

LITERATURE CITED

1883 BocouRT, Marie-Fermin. fitudes sur les Reptiles. Mission Scien-

tifique au Mexique et dans FAmerique Centrale, livr. 9, 1883, ])!. 32,

fig. 12, 12a-12c.
1940 CiLOYD, Howard K. The Rattlesnakes, Genera Sistptrus and Cro-

talm. A study in Zoogeographv and Evolution. Chicago Acad. Sci.,

Spec. Publ. No. 4, 1940, pp. vii-(-266. Index, plates, 1-31 ; text figs.

1-10; maps 1-22.
1949 Taylor, Edward H. A preliminary Account of the Herpetology of

the State of San Luis Potosi, Mexico, Univ. Kansas Sci. Bull., vol.

33, pt. 1. No. 2, Apr. 20, 1949, pp. 169-215.
19-18 Tihex Dr. J. A. A new Gerrhonotus from San Luis Potosi. Trans.

Kansas Acad. Sci.. vol. 51, 1948. ])p. 302-305.

THE UNIVEKSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. IT] March 20, 1950 [No. 12

On the Status of the Family Desmogiiathidae
(Amphibia. Caudata)

BY
E. Iv.\N SOLER.

Mayagupz, Puerto Rico

Abstract: The anatomical featuiesi of the salamanders of the genera Dcs-
mog)iathus and Leurogmathus are re-examined. Especial study of the char-
acteristics of the vertebrae is made. The author concludes that the vertebrae
differ ba>ically from tho.?e of the family Plethodontidae and that this fact,
together with numerous other anatomical differences, warrants the re-e.stab-
lishment of the family Desmognathidae. He includes the genera Lciuog-
iialhus and Desmognnthus.

INTRODUCTION

The genus Desmognathus comprises a group of common North
American salamanders, which occur from the southern extreme of
New Brunswick southward to northern Florida and southwestward
to the eastern extremes of Texas and Oklahoma. There are at
present eleven species and subspecies recognized in this genus. The
anatomy, physiology, and life history of the animals of this group
have been extensively investigated and are well known. The ready
availability of these animals where they occur makes them ideal
for such investigations. Mrs. I. W. Wilder (1913) published a com-
plete work on the life history of Desmognathus fusciis. Seelye
(1906) investigated the circulatory and respiratory systems of this
species. Fish (1895) published a detailed account of the nervous
system and its development in the same species. Other authors
also have reported upon their investigations of this and other forms
of Desmognathus. The ecology of the various species of this genus
has also been detailed at various times chiefly by G. K. Noble, E.

(459)

460 The University Science Bulletin

R. Dunn, C. H. Pope, and C. S. Brimley. A considerable literature
has appeared on the taxonomy of the salamanders of the genus
Desmognathus.

In contrast to the general agreement among workers upon the
anatomy, physiology, and life histories of the members of this
genus, the taxonomic literature indicates a marked disagreement as
to the higher taxonomic categories. Baird (1849) used this name
to denote a genus that then contained only the three known species:
D. fusais (Rafinesque) , D. auricidntus (Holbrooki [= D. fuscits
cmriculatus], and D. niger (Green) [= -D. quadramaculatus qiiad-
ramaculatus]. In 1866, Cope proposed the family Desmognathidae,
with the single genus Desmognathiis, and in 1869, he published a
complete classification of the lungless salamanders proposing an-
other family Thoriidae having a single type genus Thorivs.

Boulenger in 1882 placed the then recognized families Salaiuan-
dridae, Amblystomatidae, Plethodontidae, and Desmognathidae to-
gether in a single family Salamandridae. He arranged each of the
former families as subfamilies and placed the genus Thorius in the
subfamily Desmognathinae.

Cope again in 1889, in his "Batrachia of North America" persists
in maintaining family rank for the Desmognathids and the other
groups that Boulenger had reduced to subfamily status. Thus a
status quo was reached, with zoologists adhering either to Bou-
lenger's or Cope's classification.

In 1892 Stejneger described a new genus Typhlotriton ascribing
it to the family Desmognathidae following Cope's classification.
In 1893 Cope described a new genus Haptoglossa from Costa Rica
and revised his concept of the thoriid group reducing it to subfamily
rank as Thoriinae under the Desmognathidae and referring to it
three genera: Thorivs Cope, Typhlotriton Stejneger, and Hapto-
glossa Cope. Moore in 1899 described Levrognathus, a new genus
of the family Desmognathidae.

In 1900 Moore, basing ins opinion on iiis own observations and
those of Vaillant in 1882 and 1886, concluded that the family Des-
mognathidae was not valid. He based his conclusions solely on
the characteristics of the articulation of the vertebrae. He in-
cluded the desmognathid group in the family Plethodontidae, yet
did not exclude the possibility of its classification as a subfamily.

Fowler in 1907 and Hurter and Strecker in 1909 retained the Des-
mognathidae as a valid family. In 1908 Brown designated D. juscus
juscus (Rafinesciue) as the type species of the genus Desmognathns

Soler: The Family Desmognathidae 461

and also retained the family Desmognathidae. In Mrs. Wildcr's
work "The life history of Desmognaihus fuscus" in 1913, she con-
sidered the genus Desmognathus to be a member of the family Des-
mognathidae. However, in 1912 Brimley placed Desmognathm
as a genus of the family Plethodontidae, as did Fowler and Dunn
in 1917. Nevertheless Dunn in the same year treated the genera
Desmognathus and Leurognathus as members of the plethodontid
subfamily Dismognathinae {sic).

In 1917 Stejneger and Barbour considered Desmognathus under
the family Plethodontidae. Finally in 1926 Dunn placed Desmog-
nathus under the Plethodontidae, but did not mention his previous
subfamily Dismognathinae. Although he suggested some form of
generic grouping of the plethodonts, he made no attempt at actual
taxonomic differentiation of these generic groups. From 1926 until
the present, Dunn's classification has been generally used. How-
ever, Taylor (1944: 193 and 204) questioned Moore's reasons for
invalidating the Desmognathidae. Very recently Smith and Taylor
(1948) have come to recognize the Desmognathidae as a valid
family.

Since it has been shown that there are still uncertainties and
differences of opinion as to the higher taxonomic categories of this
group, it has become apparent that a re-examination of the problem
is absolutely necessary. Consequently I have re-examined such ma-
terial as is listed below and I have noted similarities and differences
among these groups, especially as regards the Plethodontidae. I
have tried to weigh carefully their relative importance, and on this
basis award this group of salamanders the taxonomic rank which
it seems to deserve. The following points are considered:

1. That the principle of taxonomic differentiation on tlie basis
of vertebral articulations in salamanders is still a concept that
must be applied to differentiate between the Desmognathidae and
other families.

2. That the family has other differentiating characters besides
the vertebral articulations — characters which must be taken into
consideration in evaluating the taxonomic rank and relations of
this group.

3. That the Desmognathidae is a valid family.

4. That the family Desmognathidae comprises the two genera
Desmognathus and Leurognathus. The possibility of a third genus,
Haptoglossa, is considered.

462 The University Science Bulletin

MATERIALS AND METHODS

There have been avaihable for study specimens and skeletons
of Desmognathas juscus fuscus, D. fuscus hrimleyorum, D. fuscus
auriculatus, D. ochrophaeus ochrophaeus, D. ochrophaeiis carolinen-
sis, D. quadramacidatus quadramaculatus, Leurognathus marmorata
marmorata, and L. marmorata intermedia. I have studied material
of the following plethodontid genera: Plethodc'n, Thorius, Eurijcea,
Gyrinophilus, Pseudotriton, Emantina, Chiropterotriton, Hydro-
mantes, Batrachoseps, Hemidactylium, Mancidus, Pseudoeurycea,
Magnadigita, and Bolitoglossa. I have also studied family charac-
teristics as evidenced by skeletal and preserved material of Taricha,
Diemictylus, Siren, Necturus, and Ambystoma.

The procedure followed was first, to make examination of osteo-
logical material of tlie various species of Desmognathus and Leu-
rognathus. Then comparisons were made with skeletal structures
of the genera previously mentioned. Emphasis was placed on the
morphology of the vertebral column and cranial structures although
other characters were also observed. Secondly, a comparative
study of the musclature of various species of Desmognathus and
Leurognathus were undertaken and again comparisons were made
with the other forms. Emphasis here was given chiefly to the
muscles of the head and throat.

Owing to limited time a study of the microscopic anatomy of these
forms was not feasible. However realizing the possibility that im-
portant taxonomic characters might be found in these structures, I
have endeavored to examine the more pertinent literature dealing
with the various systems and organs. From this source a consider-
able number of characters have been discerned which have direct
bearing on the problem.

ACKNOWLEDGMENTS

This work has been done under the official direction of Dr. Ray-
mond G. Stone, to whom I am grateful. I am deeply indebted to
Dr. Edward H. Taylor for the larger part of the materials used in
this stucy. I am also under obligation to the following persons
who have been most accommodating in providing me with material
otherwise not available: Dr. Sherman C. Bishop, Mr. Charles M.
Bogert, Dr. Wilmer W. Tanner, Dr. Harry G. M. Jopson, and Dr.
Robert C. Stebbins. I also extend my thanks to Mr. Lawrence F.
Smith for the preparation of the drawings here included.

Soler: The Family Desmognathidae 463

VERTEBRAL DIFFERENCES

Boulenger (1882: 2) and Cope (1889: 33) separated the Desmog-
nathidae from the Plethodontidae chiefly on the basis of their ver-
tebral articulation. The desmognathids were regarded as having
opisthocoelous vertebrae while the ]ilethodonts were considered as
being amphicoelous. Moore (1890: 621 1 considered the vertebral
articulation as the only character of significance that created dis-
tinction between these two families. He claimed that the condition
of the vertebrae is dependent on the age of individual animals, and
consequently of no value as a systematic character for the separa-
tion of families. While Hemidactylium, Plethodon, and Aneides
never become opisthocoelous, he stated that Pseudotriton ruber and
Gyrinophilus porphyriticus show a progressive ossification of the
intervertebral cartilage throughout life until old adult specimens
reach an opisthocoelous condition; and that a condition comparable
to Desmognathus is reached in P. ruber at 180 mm. and in G.
porphyriticus at 200 mm. in length. I have not been able to ob-
tain specimens for investigation with these exact measurements
In fact the largest specimen of P. ruber examined by Bishop (1946)
was only 178 mm., two mm. short of that which Moore estimates
as the probable length of his specimen. The largest specimen of
P. ruber in the Museum of Natural History of the University of
Kansas measures only 155 mm. I have procured through the kind-
ness of Doctor Bishop a specimen of G. porphyriticus 190 mm. long,
which approaches very closely the size of the specimen used by Doc-
tor Moore. I dissected both specimens and I found what might be
called a ''false" opisthocoelous condition. Immediately after dissec-
tion the anterior vertebral articulation exhibits a bulbous condyle in
both species, yet by simply touching or probing this structure one
finds out that it is soft and not ossified. This fact is further con-
firmed when the vertebra is dried out and this knob shrinks back
into the cup.

However, in Desmognathus and Leurognathus a true opisthocoe-
lous condition is achieved at an early age. I have dissected a speci-
men of D. ochrophaeus ochrophaeus of only 55 mm. and one of D.
fuscus fuscus 60 mm. in length in which already the process of ossi-
fication had extended beyond the border of the cup. Thus the
morphological results of this process of ossification are present in
Desmognathus throughout approximately the whole life of the ani-
mal. In such genera of plethodonts where this process is said to
occur, its results are present only in the very old adult stages.

464 The University Science Bulletin

Saggital sections of the vertebrae of Desmognathus, Thorhis,
Gyrinophilus (140 mm.), Pscudotriton (135 mm.), Pscudoennjcea,
Plethodon, and Taricha were made. To accomplish this the verte-
brae were imbedded in transparent plastic blocks and then the sur-
face was reduced to the desired level with fine sandpaper. The re-
sults have been decisive in demonstrating the clear cut differences in
the vertebral articulations existing between the desmognathids and
other genera of urodeles. The anterior part of the centrum of
Desmognathus, as shown in PI. X, fig. C, is a solid bony structure
filling the cup and appearing homogeneous with the walls of the cen-
trum leaving no trace of any part of a notochordal canal in the an-
terior half of the centrum of the vertebra. A solid condyle pro-
trudes anteriorly. The posterior part is typically hollow. Thus the
vertebrae are typically amphicoelous. Thorius and Taricha are
externally opisthocoelous with a well-defined condyle. However,
this is somewhat misleading since the anterior articulating condyle
is merely a convex ossified cap fitting over the anterior opening of
the cup, while the cup itself is hollow (see Pi. X, figs. A and B).

Th(> notochordal canal of the centra of Gyrinophilus and Pseudo-
triton show notochordal cartilage, but no bone. Thus for all practi-
cal purposes they are still amjihicoelous forms like Plethodon. (See
PI. X, figs. D and E). The cartilage present in the vertebrae occu-
pies only the anterior edge of the cup, leaving a hollow cavity behind
it. In still older forms this cartilage protrudes anteriorly in the
form of a condyle in which there may be some deposition of calcium.
However, this shrinks back upon drying, or if allowed to macerate
will disappear completely. In the G. porphyriticus (1!)0 nun. long),
it was found that the posterior portion of the notochordal cartilage
had become calcified or ossified, however, not filling the posterior
part of the cup. This calcified portion did not extend forward to the
edge of the cup and furthermore, unlike Desmognathus, it was easily
distinguishable from the bony walls of the centrum. The condylar
portion of the intervertebral cartilaginous mass still remained soft
and shrank upon exposure.

The term, opisthocoelous, still describes externally the vertebrae
of Desmognathus, Leurognathus, Taricha, and Thorius; and the
term amphicoelous is likewise applicable to all plethodonts except
Thorius. However, to differentiate between these forms the classifi-
cation of vertebrae now needs something more than terms descrip-
tive of external appearance. A sagittal section shows three very
distinct internal conditions of the vertebral centrum. To describe
these I pro])ose the following terms:

Soler: The Family Desmognathidae 465

The term "holocoelous," meaning entirely hollow, is proposed for
that type of vertebral centrum typical of many i^lethodoni;^ and
various other primitive salamanders. This term is descriptive of the
notochordal canal when there is no evidence of calcification or ossi-
fication of the material enclosed therein.

For the condition of the vertebral centrum in Desmognathus and
Leurognathvs the term "hemicoelous" or half hollow, is proposed.
This term is descriptive of that condition in which the posterior half
of the notochordal canal is hollow and the anterior half is solid bone
including a condyle projecting from it.

'Tseudocoelous" is proposed to describe that condition of ihe
notochordal canal such as obtains in Thorhis and Taricha. Here the
canal itself is hollow, yet it is covered at its anterior end by an ossi-
fied cap which, when observed from the outside, conveys a false im-
pression of having a condition similar to that found in Desmog-
nathus. It is also a term applicable to very old adult specimens of
Gyrinophilus and Pseudotriton where there may be calcification
within the centrum of a part of the notochordal cartilage.

But the condition of the vertebral articulation and the charac-
teristics found in the centrum are not the only differences betw^een
the genera of Desmognathidae and the genera of Plethodontidae.
Upon close examination of the vertebral column one finds further
characteristics that thus far have only been suggested (Dunn 1926,
and others), but not used in the differential taxonomy of the group.
Dunn points out that the atlas of Desmognathus and Leurognathus
is very different from that found in plethodonts. There is a gradual
curving upward of the dorsal part of the arch of the atlas w^hich
forms a sharp-angled elevation where it meets the vertically ar-
ranged supraposterior surface (see PI. XI, fig. B and C). This
strong, sharp-edged, wide elevation serves as a posterior point of
attachment for the tendon that immobilizes the temporalis muscle.
This surface formed by the elevation also serves as an anterior
point of attachment for the ends of some fibers of the dorsal longi-
tudinal muscles. The atlas of plethodonts has a plane horizontal
dorsal surface and there is no vertically arranged supraposterior
surface as such. No doubt this elevation in Desmognathus and
Leurognathus has been caused by the pulling stress of the temporal
tendon upon the dorsoposterior angle of the atlas throughout the
evolution of the forms.

The lateral articulating facets of the odontoid processes of the
atlas of most, if not all, plethodonts arc fused medially by a thin

29—90

466 The University Science Bulletin

sheet of bone. In Desmognathns and Lenrognathus this ossification
has not taken place and the two facets are distinctly separated and
very much reduced, practically nonexistent.

The remainder of the vertebral column of Desmognathns and
Leurognathus exhibits characteristics typical of most urodele verte-
brae. However, there are pointed processes arising from the dorsal
surfaces of the postzygapophyses of the anterior jM'esacral ver-
tebra which do not occur in plethodonts (see PI. XI, figs. D and E).
In younger specimens these processes are normally present only in
the first three or four presacral vertebrae, excepting the atlas, but
in older specimens these appear present to the tenth or thirteenth
vertebra. I have not observed them present posterior to the thir-
teenth vertebra.

CRANIAL CHARACTERS

In studying the skull of Desmognathns and Leurognathus, I have
followed an anteroposterior sequence. The skulls of these forms are
relatively large, yet keeping a size which is related to tlie general
proportions of the body. They are completely ossified units with
firm sutures. The anteroposterior streamlining of the skull gives
testimony to the aquatic evolution of the group. The widening of
the external surface of the spines of the premaxilla is especially
eloquent in this respect. In the characters of the premaxilla we
find the main generic difference between Desmognathus and Leurog-
nathus. The latter has no premaxillary fontanelle, while the former
has a rather large one. Nevertheless both genera have the paired
premaxillary foramina. The wedge shape of the snout in both
forms must be the resultant of the underwater "rooting" habits of
these forms.

The frontal bone that joins directly with the maxilla owing to the
absence of a prefrontal, has a ventrolaterally directed process which
arises from the side of the frontal bone, borders the anterior part of
the orbit, and forms an osseous division between the nasal chambers
and the orbits of the eyes. The origin of this process is open to
speculation. It definitely is not present in the related families
Plethodontidae and Salamandridae.

The parietals have been subjected to great stress by the temporalis
muscle and its contained tendon. This muscle runs along the dorsal
surface of the parietal where the latter forms a suture with the
periotic. It then turns ventrally on the anterior edge of the parietal
to insert on the coronoid process of the mandible. A rounded groove
in which the muscle lies, is present on the dorsal surface of the bone'

Soler: The Family Desmognathidae 467

and, furthermore, the portion of the parietal bordering the orbit has
been extended somewhat forward, thus considerably reducing the
length of the orbit. In most plethodonts the length of the orbit is
approximately one half or more of the total length of the skull.
However, in Desmognathus, owing to the condition of the parietals,
the length of the orbit is about one third of the total cranial length.

Another striking difference between the Desmognathidae and the
Plethodontidae is the morphology of the occipital condyles. In the
plethodonts which I have examined the condyles are sessile and their
articulating surface are either plane or concave. In Desmognathus
and LeurognatJms there are two stalked condyles measuring one or
two millimeters in length, extending posteriorly from the skull, and
with a convex surface to articulate W'ith the concave facets of the
atlas.

The morphology of the mandible provides a constant difference
which is undoubtedly of taxonomic value. The coronoid bone of
most plethodonts forms a low, gently tapering, elliptical curvature
on the posterodorsal edge of tiie mandible. But in Desmognathus
and Leurognathus the temporalis and its stress on the adjacent bones
have produced a change from this form, and the coronoid process
formed in these genera is an acute, pyramidal elevation of the pos-
terodorsal edge of the mandible. Its upper tip is the point of in-
sertion of the temporalis. One may readily infer that this, just as
the already described peculiar formation of the atlas and the groov-
ing of the parietal area of the skull, is a result of the stresses of the
temporal tendon. This also shows one example in amphibians of
what Gregory (1933: 446) has so painstakingly shown in fishes when
he says: "The facts cited in this paper sufficiently prove that in
the skull of the fish, just as in the mammalian skeleton, bony trabc-
culae, ridges, buttresses, etc., arise in response to specific stresses,
such as those generated by the thrusts of one moving part upon
another, in other words, that bones are usually strengthened in pro-
portion to the loads they bear."

DENTITION

Relatively little work has been done on the dentition of sala-
manders in general. The teeth of salamanders are usually very small
and in live and preserved specimens they are partially concealed
under a layer of epithelium. Nevertheless, in skeletons the denti-
tion may be easily studied under a low power microscope. Dcsmog-
nathids and plethodonts usually have pleurodont teeth arranged
along the buccal ridges of the maxilla, premaxilla, and mandible.

468 The University Science Bulletin

However there is a considerable variation among plethodonts as to
the maxilhiry teeth. While Thorius has an edentulous maxilla,*
Batrachoseps has various rows of teeth along the maxilla. Both
families also have vomerine and paravomerinef patches of teeth.
These may be arranged in either continuous or discontinuous patches.
Male desmognathids tend to lose the vomerine teeth, this effect being
more accentuated in some species than in others. This tendency to
lose completely the vomerine teeth is not found in plethodonts.

Noble in 1927 and Noble and Pope in 1929 linked dentition in
Desmognathus to secondary sexual characteristics. Secondary sex-
ual dimorphism is evident in many other urodele genera, but its
effects in desmognathids are unique and quite different from those
found in other forms. The tendency to complete loss of the vomerine
teeth seems to be restricted to desmognathids alone. Other second-
ary sexual characteristics like the elongation and forward protrusion
of the premaxillary teeth and the tendency to lose the posterior max-
illary teeth are mutually shared with some genera of plethodonts
and perhaps other families.

THE HYOBRANCHIAL APPARATUS

One of the most significant differences between the Desmogna-
thidae and the Plethodontidae is found in the hyobranchial appa-
ratus of the larvae of Desmognathus and Leurognathus. Mrs. Wilder
(1913: 300) had already noticed in her study of the life history of
Desmognathus the presence of four gill slits in the larvae of these
forms. But this condition did not seem extraordinary to her as she
correctly considered the genus to belong to the family Desmogna-
thidae. This is a radical departure from the typical condition in
the larvae of the Plethodontidae which have only three gill slits.
However, more recently, Hilton (19-1:7: 191), after examining the
hyoid apparatus, comments as follows: "In all the genera of Plethe-
dontidae examined, two only have four pairs of branchial arches,
Desmognathus and Leurognathus. This may show that these are
closely related and also suggest that they are more definitely re-
lated to other groups than the rest of the family so far known, for
they all have the rather distinctive number of three pairs of bran-
chial arches."

The adult hyobranchial skeleton of Desmognathus has been
studied by Piatt in 1935 and compared with that of various genera

* Cope in his "Batrachia of North -America," PI. 37, figs. 2-4, shows a skull labeled as
Thorius pennatulus, having maxillary teeth. This is obviou.sly an error, as all the known species
of Thorius, including T. pciinaluhts, have an edentulous maxilla.

t I follow Taylor (1914: 206) in the usage of the term iiaiaxomerine for the iiatehes of
teeth apjx'aring on two thin lion_\' plates ventral to the parasi>hi'noicl bone.

Soler: The Family Desmognathidae 469

of the Plethodontidae. The adult structures are basically similar,
except for a few details of size and shape.

MYOLOGICAL CONSIDERATIONS

Most striking muscular differences in salamanders are found in
the musculature of the neck and the throat. This general region
exhibits changes in both the superficial and the deep muscles of the
various genera which are usually associated with the adaptations
of the hyoid appartus. Desrnognathus and Leurognathus are no ex-
ceptions; in fact some of the most pertinent differentiations between
the Desmognathidae and other families are found in the character
and extent of development of this musculature.

There are two pairs of muscles in Desmognathus and Leurog-
nathus that are extraordinarily developed when compared to analo-
gous muscles in other salamanders. On the dorsal side of the ani-
mals we find the curious M. temporalis which has its origin on the
posterodorsal margin of the atlas and whose fibers encircle a strong
tendon that passes forward over the surface of the cranium, turns
ventrad on the posterior edge of the orbit of the eye and is inserted
on the tip of the coronoid process of the mandible. This tendon
immobilizes the muscle and consequently the mandible, as Dunn
(1926: 59) has indicated. To open their mouths desmognathids
have to raise the whole skull and this is accomplished by the pow-
erful dorsal and lateral muscles of the neck. Here is a condition
different from any other family of salamanders, one that I regard
as of invaluable taxonomic importance.

On the ventral side of desmognathids we find a bilobate enlarge-
ment of the gular region by means of which members of the family
can be readily recognized externally. Its prominence gives this fam-
ily what Piatt (1935: 222) has called "the characteristic Desmog-
nathus profile." This enlargement so noticeable externally, is caused
by the tremendous development of the gularis musclci in those
forms. Upon removal of the skin these muscles stand out very
prominently (see PI. XI, fig. A). They originate on the quadrate,
the articular, and the dorsolateral fascia; their insertion is on the
skin of the gular fold and the pectoral fascia.

An interesting condition in the myology of salamanders is the di-
rect correlation between the extent of attachment of the tongue and
the posterior reach of the M. subarcualis rectus, the muscular cup

t Piatt (1935: 222) and (1940: 10) has shown that this structure is really made up of two
separate nui^cles. a large quadrato-pectoralis and a small, delicate gularis. However, Eaton
(1937: 324) pointed out the convenience of using the name gularis to denominate the entire
srructure for purposes of simplicity.

470 The University Science Bulletin

that encloses the ceratobranchial cartilage and which is primarily
used in extending the tongue (Francis, 1934: 61). All desmognathids
have an attached tongue, while the Plethodontidae, with its great
variety of genera, has forms in which the tongue has become freed
from the floor of the mouth and capable of considerable protrusion.
Those plethodontid forms with attached tongues may be regarded
as more primitive. Plethodontid salamanders of the genera Eurycea
and Pseudotriton, with a tongue free all around, have a long, far
reaching subarcualis rectus, while the genera of Desmognathidae,
together with those genera of plethodonts that have the tongue at-
tached in front, have a shorter, subarcualis rectus. It seems that
the tongue attachment tends to hinder the development of the sub-
arcualis rectus, as the maximum extension of the tongue is definitely
limited to a fixed measurement by this attachment. However, in
those forms where there is no lingual attachment, the movement of
the tongue has no other limit except that imposed by the maximum
action of the subarcualis rectus itself.

Another noticeable and presumably important character is the
position assumed by preserved specimens of Desmognathus and
Leurognathus. Their head is bent downward giving a pronounced
rigor 7iiortis appearance on the anterior part of the body with the
nape raised as the apex of an inverted letter V. This would suggest
some unknown inherent quality of the musculature which is not
present in the Plethodontidae. I have not been able to observe this
peculiar condition in any other forms.

CIRCULATORY AND RESPIRATORY SYSTEMS

I have made no original study of the circulatory and respiratory
systems of the Desmognathidae. However, I have examined such
papers as would seem pertinent in this field. Among these the most
important is the article by Seelye in 1906. Here a study was under-
taken of the similarities and differences between a representative
member of the family Desmognathidae, Desmognathus fuscus, and
a member of the family Plethodontidae, Spelerpes fuscus [^Hy-
dromantes gcnei italicus\. This work was in turn partially based
on the work done on Hydromantes by Betiige in 1898. No differ-
ences were found in the respiratory system, yet a resume was made
of the differences in the circulatory system which I quote as follows:

"1. The vena subclavia entering the sinus venosus directly, in-
stead of first opening into the ductus Cuvieri; and the right and left
venae subclaviae opening together into the sinus on its left side.

Soler: The Family Desmognathidae 471

"2. Tlie vena abdominalis arising, not from the venae iliacae, but
from the venae iliacae communes.

"3. A side tributary of the vena lingualis, observed, however, only
on the left side.

"4. The arteria cutanea, a branch of the arteria pulmonalis, in-
stead of a branch of the arteria subclavia.

"5. Four arteriae gastricae at anterior end of stomach, instead of
two; three of these go to one side (the left).

"6. The presence of the arteria epigastrica, an important branch
from the aorta just anterior to the arteria iliaca."

After considering these facts one is led to the conclusion that these
differences are of taxonomic importance. It is not only one single
detail, but an accumulation of differences between Desmognathus
and Hydro7nantes in the morphological structures of one single sys-
tem which seem to indicate no less than distinction on the family
level.

REPRODUCTIVE SYSTEM

The testes and vasa deferentia of all members of the family Des-
mognathidae are covered by a deeply pigmented mesentery of coal
black color. Since the first part of the century many biologists have
tried to associate testicular pigmentation with external stimuli.
Considerable evidence has been accumulated on the effects of light
and temperature upon this pigment, but most of it is conflicting.
Only of late, Finkel in 1945, has finally and definitely linked tes-
ticular pigmentation with sex hormones in opossums and ground
squirrels. This work does not eliminate the strong possibility of
establishing a correlation with external stimuli, but on the contrary
it has indicated the necessity for such further investigations.

A few members of the genus Plethodon of the family Plethodon-
tidae likewise have developed pigmented genitalia. However, I be-
lieve this case is not an illustration of relation, but one of parallel
development.

NERVOUS SYSTEM

In the literature of the nervous system of salamanders, Murray
(1897) has made a comparison between the ramifications of the
spinal chord and nerves of the three forms: Desmognathus fuscus,
Plethodon erythronotus [= P. dorsalis], and Spelerpes fuscus
[=^Hydromantes genei italicus]. He has noted a characteristic
which I believe adds weight to the differentiation of the families
Desmognathidae and Plethodontidae. He found that in the three
forms the spinal nerves pierce the neural arch, however in Plethodon

472 The University Science Bulletin

the dorsal and ventral roots of the spinal nerve are separated by
a ridge of bone on the inner surface of the neural arch. The fora-
men of exit, however, is single. A different condition from this
last one obtains in Desviognathus where the dorsal and ventral roots
of the spinal ner\-es leave the osseous canal by separate foramina.

GLANDS

Another characteristic of the soft anatomy of the Desmognathidae
which undoubtedly has bearing on taxonomic differentiation between
these forms and plethodonts, is that which Mrs. Wilder (1913: 315)
has noted when she studied the development of the naso-optical
region in Desviognathus. I ciuote: 'Tn connection with the develop-
ment of the orbital glands it should here be noted that unlike all
Urodeles (so far as the facts have been reported), Desmognathns
has no nasolacrimal ducts. As these are present in so closely re-
lated and associated forms as Spelerpes and Plethodon, as well as
in the less closely related lunged forms, their absence here has some
significance which demands further study of the comparative mor-
phology and the habits of these species."

The members of the Desmognathidae develop a submental gland
in the male as a secondary sexual characteristic. Dunn (1926) and
Noble (1927) have already discussed these glands. Their function
seems to be hedonic. In the Desmognathidae these glands are re-
stricted to the very anterior tip of the chin, immediately ventral to
the mandibular symphysis. In this family they are composed of
five or six separate fingerlike glandules arranged in a typical shape.
There are submental glands in the Plethodontidae; however, their
characteristics are very different from the analogous structures in
the Desmognathidae. In the genera of plethodonts that I have ex-
amined the glands are flat, circular clusters of globules attached to
the inner side of the skin just ventral to the floor of the mouth.

There are three sets of glands located along the walls of the
cloaca of male salamanders. These glands have been found by
Noble and Pope (1929) to have reproductive and hedonic functions.
The anterior pair are the pelvic glands whose secretion mixes with
the spermatozoa to form the head of the spermatophores. The
l^iapillae of the cloacal glands form the greater part of the villous
pad on either side of the cloaca. The abdominal glands simulate a
pair of wings on the posterior angle of the cloaca. In the Desmog-
nathidae the abdominal glands have a peculiarly hard texture when
compared to the same glands of other families and to the softer and
fluffier pelvic and cloacal glands of the same forms. In many speci-

Soler: The Family Desmognathidae 473

mens I have observed a peculiar coloration of these abdominal
glands that varies from light pink to light brown. I have not found
this coloration in any species belonging to other genera.

CHARACTERS OF THE FAMH.Y DESMOGNATHIDAE

Future work on this problem, if directed to the soft anatomy, and
especially the microscopic anatomy of the Desmognathidae will
doubtless yield added evidence for the differentiation of the Des-
mognathidae from the Plethodontidae. Points of resemblance be-
tween the two families suggest common ancestry, or parallel devel-
opment. The facts evinced from this re-examination of the problem
more than suffice to confirm the separation into the two distinct
families. In summation, the following combination of characters
may be regarded as defining the family Desmognathidae:

1. True opisthocoelous vertebrae.

2. Hemicoelous centra.

3. A nasolabial groove present.

4. Lungs absent.

5. Tongue adherent in front and by the middle.

6. Larvae with four epibranchials.

7. Highly modified atlas.

8. Carpus and tarsus not ossified.

9. Strong tendon present in j\I. temporalis which immobilizes
the mandible.

10. Highly developed gularis muscles.

GENERA OF DESMOGNATHIDAE

The family Desmognathidae, as known today, comprises the two
genera Desniognathtis and Lenrognathus. They are almost identical
when observed externally. The main external difference between
them is the presence of a groove from the eye to the angle of the
jaw in Des7nognathus and its absence in Leurognathus. This near
identity has resulted in confusion and Leurognathus has often been
mistaken for Desmognathus by workers in the field. However,
Moore (1899), Bishop (1924), and Pope (1924 and 1928) have noted
the behavior peculiar to each of these genera and on this basis they
are able to distinguish them. When Leurognathus is exposed in
water by turning a stone it either lies quietly on the bottom or
glides gracefully to cover under water. This response is in sharp
contrast to the darting, vigorous swimming movements of Desmog-
nathus quadramaculatus or Desmognathus phoca when disturbed.

The basic generic differences between Desiyiognathus and Leurog-

474 The University Science Bulletin

nathns are easily discerned upon examination of the skull. Neither
the prcmaxilla nor the vomerine bones of Leurognathus embrace a
fontanelle as in Desmognathus. This condition gives the anterior
part of the skull of Leurognathus a completely roofed appearance.
The choanae of Leurognathus are small foramina hidden in a fold
at the sides of the roof of the mouth. In Desmognathus, however,
these arc conspicuous, plainly observable openings. The ratio of
the length of the orbit to the total length of the skull is higher in
Leurognathus than in Desmognathus. Nevertheless after these
generic differences are noticed we find that endless stream of simi-
larities which classify these two forms as two closely related genera
within the family Desmognathidac.

The loss of the type of the mysterious Haptoglossa pi'essicauda
Cope allows us only to speculate as to the kinship of this form. This
species was placed by Cope in the subfamily Thoriinae under the
family Desmognathidac. From the description given by Cope
(1893: 334) this genus could belong to the family Desmognathidac
as we see it today. However, we shall never determine its exact
taxonomic position until the species is again found. Dunn (1926:
429) doubtfully synonymizes this form with Oedipus uniformis
[= Oedipina uniformis], yet I fail to understand how he can recon-
cile the adherent tongue, the compressed tail, and the relative sizes
of the head, body, and tail as described by Cope with those of
Oedipina uniformis that has totally different characters. This form
could also be a plethodont, as none of its characters is conflicting
with the wide variety found in this family. The family association
will depend upon ascertaining the character of the vertebral struc-
ture and articulation, and other family characteristics.

As previously stated Thorius Cope and Typhlotriton Stejneger
were at one time included in the family Desmognathidae. Thonus,
however, may be promptly eliminated from it on the basis of its
pseudococlous centra, dorsal skull characters, the orbitolabial groove,
the normal atlas, its free tongue, and the absence of highly developed
gular muscles. I have not examined adult specimens of Typhlotri-
ton, but from Moore (1900: 620) I infer that it has pseudococlous
vertebrae. This last genus may be also eliminated from the family
Desmognathidae on the strength of its normal atlas and the presence
of a prefrontal bone. Dunn (1926: 251) has conclusively associated
this genus with the plethodontid group close to Eurycea, and proved
its lack of relationship with the family Desmognathidae.

Soler: The Family Desmognathidae 475

LITERATURE CITED
Baird, S. F.

1849. Revision of the North America Tailed Batrachians, with descriptions
of new genera and species. Jour. Acad. Nat. Sci. Philadelphia, 1: '
281-292.

Bethge, E.

1898. Das blutgefasssystem vom Salamandra moculata, Triton taeniatus,
und Spelerpes fuscus; mit betrachtungen liber den ort der athmung
beim lungelosen Spelerpes fuscus. Zeits. f. wiss. Zool., 63: 680-707.

Bishop, S. C.

1924. Notes on Salamanders. 19th Report of the Director. New York

State Mus. Bull., pp. 87-96.
1947. Handbook of Salamanders. The Salamanders of the United States,

of Canada, and of Lower California. 555 pp., Comstock Publishing

Co., Ithaca, New York.

BotLErN'GER, G. A.

1882. Catalogue of the Batrachia Gradientia s. Caudata and Batrachia
Apoda in the collection of the British Museum. Taylor and Francis,
London. 124 pp., 9 pis.

Brimley, C. S.

1912. Notes on the Salamanders of the North Carolina mountains with de-
scriptions of two now forms. Proc. Biol. Soc. Washington. 25:
135-140.

Brown, A. E.

1908. Generic types of Nearctic Reptiles and Amphibia. Proc. Acad. Nat.
Sci. Philadelphia, pp. 112-117.

Cope, E. D.

1866. On the structures and distribution of genera of the arciferous Anura.

Jour. Acad. Nat. Sci. Philadelphia. 6: 67-112.
1869. A review of the species of the Plethodontidae and Desmognathidae.

Proc. Acad. Nat. Sci. Philadelphia, pp. 93-118.
1889. The Batrachia of North America. Bull. U. S. Nat. Mus , No. 34.

495 pp., 86 pis.
1S93. Second addition to the knowledge of the Batrachia and Reptilia of

Co.sta Rica. Proc. Ame.-. Philos, Soc, 31: 333-347.

Dunn, E. R.

1917. The Salamanders of the Genera Desmoqnathus and Lcuroqiiathus.

Proc. U. S. Nat. Mus., 53:393-433.
1926. The Salamanders of the Family Plethodontidae. Smith College

50th Anniversary Publications. 441 pp.

Eaton, T. H., Jr.

1937. The gvilaris muscle in Urodela. Jour. Morph.. 60: 317-324.

FiNKEL, M. P.

1945. The Relation of Sex Hormones to Pigmentation and to Testis
Descent in the Opossum and Ground Squirrel. Amer. Jour. Anat.,
76: 93-151.

Fish, P. A.

1895. The Central Nervous System of Desmognathus fusca. Jour. Morph.
10: 231-286, pis. 13-16.

Fowler, H. W.

1907. The Amphibians and Reptiles of New Jersey. Annual Report of
the New Jersey State Museum, pp. 23.

FoAVLER, H. W., and Dunn, E. R.

1917. Notes on Salamanders, Proc. Acad. Nat. Sci. Philadelphia. 69: 7-28.

Fr.^ncis, E. T. B.

1934. The Anatomy of the Salamander. Oxford University Press. 376
pp., 25 pis.

476 The University Science Bulletin

Gregory, W. K.

1933. Fish SkulLs: A study of the Evohition of Natural Mechanisms.
Trans. Amer. Philos. See, 23: 75-481.

Hilton, W. A.

1947. The Hvobranchial Skeleton of Plethodontidae. Herpetologica.
3: 191-194.

HuRTER J., and Strecker, J. K.

1909. Amphibians and Reptiles of Arkaasas. Trans. Acad. Sci., St. Louis,
18: 11-27.

Moore, P. J.

1899. Lcurugnathus marmoratn. a new genus and species of salamander
of the family Desmognathitlae. Proc. Acad. Nat. Sci. PhiladeJpliia,
l)p. 316-323, 'l pi.

1900. Post-laryal Changes in the yertebral articulations of Spelerpes and
other salamanders. Proc. Acad. Nat. Sci. Philadelphia, pp. 613-622.

Murray, J. A.

1897. The Vertebral Column of certain primitive Urodela : Spelerpes,
Plethodon, Desmognathus. Anat. Anz., 13: 661-664.

Noble, G. K.

1927. The Plethodontid Salamanders; Some Aspects of their Eyolution.
Amer. Mus. Novit. No. 249, 26 pp.

Noble, G. K., and Pope, S. H.

1929. The Modification of the Cloaca and Teeth of the Adult Salamander,
Dcsniognathus, by Testicular Transplants and by Castration. British
Jour, of Exp. Biol., 6: 399-411, pis. 8-9.

Piatt, J.

1935. A comparatiye study of the hyobranchial apparatus and throat
musculature in the Plethodontidae. Jour. Morph., 57: 213-242, 4 pis.

1940. Correct Terminology in Salamander Myology. II. Transverse
Ventral Throat Musculature. Copeia, pp. 9-14.

Pope, C. H.

1924. Notes on North Carolina Salamanders with Especial Reference to
1 the Egg-laying Habits of Leurognathus and Desmognathus. Amer.

Mus. Novit. No. 153. 15 pp.

1928. Some Plethodontid Salamandeis from North Carolina and Kentucky
with the Description of a New Race of Leurognathus. Amer. Mus.
Novit. No. 306, 19 pp.

See lye, A. B.

1906. Circulatory and respiratory systems of Desmognathus fusca. Proc.
Boston Soc. Nat. Hist., 32: 335-357.

Smith, H. M., and T.^ylor, E. H.

1948. An Annotated Checklist and Key to the Amphibia of Mexico.
U. S. Nat. Mus. Bidl. 194, 118 pp. '

Ste.jxeger, L.

1892. Preliminary Description of a New Genus and Species of Blind Cave
Salamander from North America. Proc. U. S. Nat. Mus., 15: 115-117.

Stejneger, L., and Barbour, T.

1917. A Check List of North American Anii)hibians and Ro]>tilcs. 125
pp.. Harvard University Press.

Taylor, E. H.

1944. The Genera of Plrthodont Salamanders in Mexico. Part I. Univ.
Kan.sas Sci. Bull.. 30: 189-232.

Soler: The Family Desmognathidae 477

V.MLLAXT, L.

1882. Remarque sur la disposition des corps vertebraux chcz VAne'ides
lugubris Hallowell. Bull. Soc. Philomathique de Paris, 6: 183.

1886. Note complementaire sur I'anatomie de VAne'ides lugubris Hallo-
\v(>ll. 2d Bull. Soc. Philomathique de Paris. 10: 42-43.

Wilder. I. W. [Mrs.]

1913. The life history of Dcsmogiuithus fusca. Biol. Bull., 24: 251-343.

478 The University Science Bulletin

EXPLANATION OF PLATE X

Fig. a. Thorius norisovalis, No. 25216 EHT-HMS.

Fig. B. Tancha torosa, No. 18263 M.V.Z.K.U.

Fig. C. Desmognathus fuscm fuscus, No. 28293 EHT-HMS.

Fig. D. Pseudotriton ruber ruber, No. 19450 M.V.Z.K.U. (.specimen 135
mm. in length).

Fig. E. Gyrinophilus porphyriticus porphyriticus, No. 28272 EMT-HMS
(siiecimen 140 mm. in length).

PLATE X

Sagittal sections of presacral vertebrae (enlarged)

480

The University Science Bulletin
PLATE XI

riiJi B

■tt^^

\ ) c

Fig. a. Ventral view of the throat musculature of an adult male of Dc.s-
mognathus ochrnphaeus orhmphaeus, .^howinj: the development of the gularis
mu.scles and the submental sland.

Fig. B. Dorsal view of th(> atla.s of Desmoc/iidthus juscus fuscns showing
the reduction of the odontoid process.

Fi(!. C. Side view of the atlas of D. fuscus fuxrits showing the transverse
ridge for the attachment of the temporal tendon.

Fig. D. Posterior end view of anterior presacral vertebra of D. fuscus
fuscus showing the processes arising from the postzygaphophysis.

Fig. E. Side view of D.

THE UNIVEfiSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. II] March 20, 1950 [No. 13

Ceylonese Lizards of the Family Scincidae

BY

Edward H. Ta-v-lor,

Department of Zoology, University of Kansas

Abstr-act: This paper treats of Ceylon scincoid lizards in the collections
of the United States National Museum, together with those in the Edward
H. Taylor-Hobart M. Smith Collection at Lawrence, Kansas.

In this paper the following species are described as new: Mabuya floxceri,
Riopa dngha, Sphenomorphus deignani, Sphenomorphus rufogulus, and Nessia
deraniyagalai. One species, Mabuya madardszi Mehely is removed from the
synonymy of Mabuya macularia Blyth, and re-established.

All the Ivnown forms are listed and keys are given for the species of the
various genera. Altogether six genera occur with the 25 species distributed
as follows: Mabuya, 6 species; Dasia, 1; Riopa, 2; Sphenomorphus, 7; Chal-
cidoseps, 1 ; and Nessia, 8.

This is the second * of a series of studies on the herpetological
fauna of Ceylon. The materials on which this study has been based
are the Ceylonese specimens of the U. S. National Museum and those
in the Edward H. Taylor-Hobart M. Smith collection at Lawrence,
Kansas (EHT-HMS). In the first collection there are 367 speci-
mens; in the latter, approximately 1,000.

The source of the material in the U. S. National Museum is
varied. Certain specimens represent the result of exchanges with
various European museums; some w^ith the museum in Colombo,
Ceylon. A considerable number are from the J. Hurter collection
that likewise contained material from a variety of sources. Nearly
one third of the collection is a recent lot collected by Dr. H. G.
Deignan of the National Museum, in the Kandy and Matale Dis-
tricts of the central highland region of Ceylon.

* Edward H. Taylor. Comments on Ceylonese Snakes of the Genus Typhlops with descrip-
tions of new species. Univ. Kansas Sci. Bull., vol. 31, pt. 2, Nov. 1, 1947, pp. 283-298 fi^s
1-3. (First study.)

(481)

30—90

482

The University Science Bulletin

The material in the EHT-HMS collection, with a few exceptions,
is a recently collected lot obtained by me, chiefly from the region
about Trincomalee, Ceylon, with a scattering of specimens from
other localities on the island.

The present study deals with the lizard family Scincidae, which is
very well represented by six genera and 25 species. Five of this
lot are regarded as new, and one form, Mabuya madardszi Mehely,
is removed from the synonymy of Mabuya macularia and re-estab-
lished.

The following forms occur in Ceylon:

* Mabuya bibronii

* Mabuya macularia

* Mabuya beddomii
Mabuya floweri

* Mabuya carinata
Mabuya madaraszi

* Riopa punctata
Riopa singha
Dasia haliana

* Sphenomorphus dussumieri
Sphenomorphus megalops
Sphenomorphus deignani
Sphenomorphus striatopunctatus

Sphenomorphus taprobanense
Sphenomorphus fallax
Sphenomorphus rufogulus
Chalcidoseps thwaitesi
Nessia burtonii
Nessia didactyla
Nessia monodactyla
Nessia bipes
Nessia sarasinorum
Nessia hikanala
Nessia layardi
Nessia deraniyagalai

The faunas of Ceylon, as regards their relationship to those in
nearby land masses, seem to be paradoxical at times. Thus in cer-
tain genera the relationship is close; in others there is no apparent
relationship whatever. Thus Mabuya is represented by six species
in Ceylon, four of which are regarded as identical with those in
India; two are endemic, or at least have not yet been reported in
mainland India. On the other hand, Nessia has eight species, none
of which occurs in India. In this case the relationship seems to be
with South Africa and Madagascar where the genus Acontias occu-
pies the same niche and has undergone a somewhat similar evolu-
tion. Most, if not all, of its African and Madagascar species have
lost the limbs, and the scales are more highly specialized. Boulenger
(1887) t regarded Acontias and Nessia as belonging to a single
genus. However, a more recent proposal has been made for their
separation by Hewitt (1929)$.

* Species marked with an asterisk represent forms also occurring in India.
t Cat. Lizards British Mus., vol. 3, 1887, p. 424.
t Ann. Transvaal Museum vol. 13, 1929, p. 8.

Taylor: Ceylonese Lizards 483

Essex (1928)11 has suggested that Acontias and Nessia represent
the same stock but that they are end products of two divergent
streams from an unknown center. That this center is not in continen-
tal Asia seems obvious.

Chalcidoseps is a relict monotypic genus known only from Ceylon,
and may represent a survivor from a group, similar to Nessia, that
has undergone a similar evolution. There are other monotypic
genera of the same sort scattered throughout the southeastern part
of Asia and the Indo-Malayan Archipelago. They likewise seem-
ingly represent survivors of former specialized series of species.
Barkudia and Sepsophis in India are typical examples. I am con-
vinced that Lygosoma quadrupes § represents a single remaining
form of a similarly specialized group. I would consider Ateucho-
saurus as a representative of another specialized group; Ophioscin-
cus represents another series with three living forms.

The two species of Riopa are closely related {singha) , or identical
(punctata), wdth Indian species. Dasia, belonging to an arboreal
group of skinks, seems to be related most closely to species that
exist in the Indo-Malayan Archipelago and the western Pacific Is-
lands. A south Indian form exists (subcoerulea) , but the relation-
ship of this species is with eastern Asiatic forms and not with the
Ceylonese species, despite the fact that there is a very considerable
hiatus in the distribution of the genus in central and eastern India,
Burma and Siam.

Mabuya is a genus of wide distribution (South Asia, Indo-Ma-
laya, Africa, Central America, South America and West Indies).
It is ultraconservative and, like Eumeces, has developed no very
highly specialized types, although there is a tendency to greater
specialization (small size) in Florida and west Mexico in the latter
genus. In ultraconservative species, changes of lesser extent are
perhaps of greater significance than in the more plastic groups.

I Studies in Reptilian Degeneration. Proc. Zool. Soc. London, (1927) 1928, pp. 879-945,
text figs, and plate.

§ I cannot agree with Dr. Malcolm Smith, 1937, in considering Lygosoma quadrupes as one
of a specializing or "degenerating" series that includes Sphenomorphus and perhaps other genera
(see Rec. Indian Mus., vol. XXXIX, part III, 1937, pp. 213-234).

484 The University Science Bulletin

Key to the Genera of Scincidae in Ceylon

1. Palatine bones in contact or overlapping along medial palatal line 2

Palatine bones not in contact along medial palatal line 6

2. Supranasals present 3

Supranasals absent 5

3. Scales keeled with from 2 to 7 keels 4

Scales smooth ; body somewhat elongated ; limbs pentadactyl, but short ; widely

separated when adpressed ; scales smooth ; lower eyelid with a semitransparent
disk Riopa

4. Pterygoid bones not in contact, the palatal notch extending forward to level of

centers of eyes ; limbs pentadactyl, well developed, distinctly overlapping when

adpressed ; no distinct transverse bands Mabuya

Pterygoid bones in contact anteriorly, the palatal notch not reaching forward to
level of centers of eyes. Limbs pentadactyl, failing to touch or barely over-
lapping when adpressed; color pattern of transverse bands Dasia

5. Limbs pentadactyl, meeting, overlapping or failing to meet when adpressed ; lower

eyelid scaly; frontoparietal single or double; scales smooth or at most, with

slight suggestion of keels or striations Sphenomorphxis

Limbs not pentadactyl, greatly reduced or absent ; body anguiform ; the nostril in
rostral 0

6. Nostril in anterior part of rostral, connected to the posterior edge of scale by a

groove ; limbs variable, never tetradactyl Nessia

Nostril near the posterior edge of rostral; limbs short, tetradactyl CJialcidoseps

Genus Mabuya Rafinesque

Mabuya Rafinesque, Anal. Nat., 1815, p. 76 n.n.

Genotype. — Lacertus mabouya Lacepede (not Scincus Sloanii
Daudin fide Malcolm Smith, Fauna of British India, Rept. Amph.,
vol. 2, Sam-ia, 1935, p. 257).

Six species are recognized as occurring in Ceylon.

Key to Species of Mabuya in Ceylon

1. A transparent disc on lower eyelid bibronii

No transparent disc, but several larger transparent scales on lower eyelid 2

2. A postnasal ynacularia

No postnasal 3

3. Scales almost smooth or with three to five very feeble keels beddomii

Scales not smooth, each bearing 3, 5, or 7 strong keels 4

4. Scales tricarinate (or sometimes also with a small tubercle on outer edge of scale) ;

dorsum with a paired series of black markings floweri

Scales with five or seven keels (except very young which may have three) ; no
paired series of black markings on dorsum 5

5. Larger; snout to vent, 125 mm. ; adpressed hind limb to wrist or elbow carinata

Smaller; snout to vent, 77 mm. ; adpressed hind limb to axilla or farther. . . madardszi

Mabuya bibronii (Gray)

Tiliqua bibronii Gray, Ann. Mag. Nat. Hist., vol. 2, Dec, 1838, p. 290 (type locality un-
known).

Mabuia bibronii Haly, Report on the collection of Reptiles and Batrachia in the Colombo
Museum; Colombo, 1891, p. 14 (Mullaittivu, Eastern Province); Annandale, Spolia Zeylanica,
vol. 3, pt. 11, Jan., 1906, p. 190 (Ceylon); Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 16,
pt. 2, 1931, p. 164; Hora, Rec. Indian Mus., vol. 29, pt. 1, p. 1, pi. 1, fig. 4 (comment on
habitat).

Mabuya bibronii Smith, The Fauna of British India including Ceylon and Burma; Reptilia
and Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp. 260-261 (Chundikulam, Eastern Province).

No specimens of this species are at hand. Its known distribution
in Ceylon seems to be on the eastern coast.

Taylor: Ceylonese Lizards 485

It may be distinguished from other known Ceylonese forms by the
presence of a rounded, undivided, semitransparent disk in the lower
eyelid; two pairs of nuchals; scales 5-to 7-keeled; a postnasal, and
28-30 scale rows. The snout to vent length is 50 mm. It has been
reported only from Eastern Province.

Mabuya macularia (Blyth)

Euprcpes macularius Blyth, Journ. Asiatic Soc. Bengal, vol. 22, 1853, p. 652 (type locality,
?Rangpur, Bengal).

Mabuia macularia Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 16, pt. 2, 1931, p. 165.

Mabvya macularia (part) Smith, Fauna of British India including Ceylon and Burma;
Reptilia and Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp. 264-266.

The collection contains two specimens of this species (U. S. N. M.
No. 29410 and 120325) collected by D. L. Karcher, Ceylon, Clodagh
Estate, Rattota, Matale District. These differ in certain charac-
ters and if the differences are not due to age, may possibly represent
two forms.

No. 29410. Part of rostral visible above less than half length of
frontonasal which it touches, separating the internasals; latter
nearly one third wider than long; prefrontals forming suture; parie-
tals separated by interparietal which is distinctly longer than broad ;
two supraoculars touch frontal; length of parietal l^/s times in
width; parietal partly segmented on left side; a postnasal present;
first loreal at least twice as high as long, about half length of pos-
terior loreal; four supraoculars; five superciliaries ; three anterior
temporals; lower eyelid with quadrangular scales; seven upper la-
bials ; mental much shorter than postmental ; first pair of chinshields
separated; second pair separated by a single scale; third pair (one
of the pair segmented longitudinally) separated by three scales;
seven (or eight) lower labials; 25 scale rows, the 12 ventrals
smooth, the remainder keeled with five, six, or seven keels; tem-
porals practically smooth; scales under base of tail small be-
coming widened posteriorly both ventrally and dorsally; subdigital
lamellae on fourth toe, 13-15. Eight or ten axillary rows of granular
scales, forming a pocket; two small granular rows behind leg inser-
tion; no modified scales on posterior surface of thigh; hind leg
reaches somewhat beyond wrist of adpressed arm; nearly uniform
olive above with a broad lateral brown band bearing numerous
streaks or flecks of white.

No. 120325 is a larger adult that differs from the smaller (and
younger) in several points: The prefrontals separated, and first
supraorbital separated from frontal; series of scales in the axilla
somewhat fewer and pocket not well defined ; scales on posterior part

486 The University Science Bulletin

of femur show distinct modification in having a small patch of en-
larged soft scales near limb insertion.

There is of course the possibility that two separable forms are
represented by these two specimens. They agree in the following
characters: The first chinshields are separated, and the mental is
much narrrower than the postmental. The part of the rostral visi-
ble above is less than half the length of the frontoparietal. The
white spots on the sides are strongly evident. The hind limb
reaches half way between wrist and elbow of the adpressed arm.

A larger series of specimens will be necessary to determine the
status of the variation.

Mabuya beddomii (Jerdon)

Euprepes beddomii Jerdon, Proc. Asiatic Soc. Bengal, Mar., 1870, p. 73 (type locality, My-
sore, India).

Mabuya beddomii Smith, The Fauna of British India including Ceylon and Burma. Rep-
tilia and Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp. 274-275 (Punduloya).

A series of four specimens taken 12 mi. N of Trincomalee are
referred to this Asiatic species.

The characteristics of these four specimens follow: supranasals
broadly in contact; frontonasal very much reduced, much wider
than long; prefrontal broadly in contact; six superciliaries ; a pair
of nuchals; no postnasal; eyelid with three or four enlarged scales;
first pair of chinshields separated; dorsal and lateral scales rather
weakly keeled, with three or (rarely) five keels; temporal scales
smooth; the length of parietal contained in width, 1^ times; height
of subocular about one half its length; 30 or 31 scale rows around
middle of body.

Dorsal surface brown with a short, whitish, black-edged stripe
beginning on nuchals and extending to a line drawn in front of arm
insertion; a broad dorsolateral dark stripe to some distance on tail
bordered above by a very narrow cream line and below by a broader
(two half scale rows wide) white or cream line; this light line bor-
dered below by a dim dark line ; lateral cream line, arising on upper
lip passes for the most part above, rather than through, ear; venter
immaculate white.

These Ceylon specimens differ from the continental Indian speci-
mens in having the middorsal line much reduced. The very young
(26 mm.) specimens show the same marking as obtains in the largest
(53 mm.). Whether there are likewise stable differences in squa-
mation cannot be determined from the material at hand.

Taylor: Ceylonese Lizards 487

Mabuya floweri sp. nov.

Type. EHT-HMS No. 30507. Collected 12 mi. north Trinco-
malee, Ceylon, Sept. 3, 1944. E. H. Taylor, collector.
Paratype. No. 30506, same locality and collector.

Diagnosis. A small, terrestrial species; scales for the most part
tricarinate having transparent part of eyelid broken into three or
four short quadrangular scales; frontal short, equal to or minutely
less than its distance from tip of snout, shorter than combined
length of interparietal and frontoparietals; no postnasal; supra-
nasals separated; prefrontals and parietals also separated; first
chinshields separated; six superciliaries; a well-defined, swollen
(glandular?) area on posterior part of thigh, covered with large,
irregular, more or less pointed scales; 30 scale rows about body,
the 12 ventral scale rows smooth, the 18 dorsal and lateral scale
rows heavily keeled.

Description of the type. Rostral folding back over snout, the
part visible above less than half length of frontonasal with which
it is in contact, thus separating supranasals; latter scales narrow,
elongate, separated by a distance equal to one third of their length;
frontoparietal nearly one fifth wider than long, forming a suture
with frontal and separating large prefrontals; frontal about one
fifth longer than frontoparietals ; interparietals short, about as wide
as long; greatest length of parietals contained in width about one
and one fourth times; a well-developed pair of nuchals; second
supraocular alone touching frontal, both second and first touching
prefrontal ; four supraoculars ; suture between rostral and first labial
when extended upward passes somewhat in front of nostril; nasal
definitely a single scale; no postnasal present; anterior loreal dis-
tinctly higher and narrower than second; two presuboculars ; four
postsuboculars diminishing in size, the upper separated from last
supraocular by a single scale that is in contact with last super-
ciliary; three primary and three secondary temporals; four supra-
labials precede and two follow the subocular which is two and one
half times longer than high ; three bluntly pointed ear lobules ; four
large rectangular scales on eye; eight or nine infralabials; a large
postmental in contact posteriorly with a small median scale sepa-
rating the first pair of chinshields ; second pair separated by a single
scale, third pair by three scales; ear opening circular, its diameter
less than transparent part of eyelid; scales in 32 rows about body,
12 ventral rows smooth, 18 dorsal and lateral rows strongly tri-

488 The University Science Bulletin

carinate with sometimes an additional outer tubercle or keel on
scales from nuchals to base of tail; the keels form nearly continuous
lines arranged in groups of three, these groups distinctly separated
from the next series of three by a space that is distinctly wider than
that between keels of the rows forming the series; dorsolaterally
the outer edges of the scales show a tiny tubercular elevation or a
fairly well-defined keel; on tail, scales are bi- or tricarinate; nuchals
and scales immediately following nuchals have four to six keels;
the crease of the axilla with five or six rows of minute granules not
forming a "pocket" or at most only a very shallow one; in crease
behind insertion of thigh a similar series of five or six rows of
granular scales. Scales on arm tricarinate and either bi- or tri-
carinate on upper surfaces of thigh and tibia ; a pair of distinctively
enlarged preanals with three other smaller preanals on each side;
temporal scales keeled.

Tubercular scales on palm and soles somewhat pyramidal; sub-
digital lamellae rounded, nearly smooth without keel, 15-16 under
fourth toe. Swollen area on posterior part of thighs covered with a
series of irregular enlarged scales with narrowed tips that stand
nearly erect and are separated from each other; the scales above
and below them are regularly imbricating scales. Adpressed hind
limb reaches to elbow of adpressed arm.

Color. Above dull olive with metallic reflections ; a greenish white
dorsolateral line runs from eye to base of tail becoming less distinct
as tail is approached; a broad brownish band follows along side
from eye to groin; an indistinct line on supralabials becomes dis-
tinct behind eye, and gives a whitish border to ear opening and
beyond which it may be traced below the brown band to groin.
Top of head brownish; two series of about twenty short narrow
transverse black marks beginning at shoulder continue back to some
distance on tail. Ventral surface light with a slight greenish cast.

Measurements in mm. of the type and paratype respectively.
Snout to vent, 56, 55; tail length, 95, (?) ; length of head to end of
interparietal, 10, 9.9; width of head at ear, 9.5, 9.6; snout to ear
opening, 11.7, 11; snout to foreleg, 20, 20; axilla to groin, 27, 28;
foreleg, 14, 14.5; hind leg, 21, 21.5.

Variation. The paratype agrees with the type in all characters
listed except there is less evidence of an outer keel in the smooth
space between the series of three keels, and the small dorsal marks
are reduced to two dots instead of a single narrow transverse mark.
These double series occur from shoulder to some distance on the tail.

Taylor: Ceylonese Lizards 489

Remarks. This species was found in a coconut grove some distance
back from the sea beach and was never observed climbing.

The modified scales on the posterior part of the thigh is a character
that has not been used by Boulenger or Smith in separating the
Asiatic species of Mabuya but it is apparently constant in the species
in which it occurs*. This condition is reminiscent of a scale modi-
fication that occurs on the posterior part of the thighs of certain
eastern Asiatic Eumeces.

The looseness of these scales allows the ingress of numerous small
mites, and both specimens show infestations of these parasites.

Mabuya carinata (Schneider)

Scincus carinatus (part) Schneider, Historia Amphibiorum, vol. 2, 1801, p. 183 (type locality
unknown).

Mabuia carinata Boulenger, Catalogue of the Lizards in the British Museum, 2d ed., vol. 3,
1887, pp. 182-183 (Ceylon); Fauna of British India including India and Ceylon; Reptilia and
Batrachia, 1890, pp. 188-189, fig. 56- (Ceylon). Willey, Spolia Zeylanica, vol. 4, 1907, pp. 186-
188. Green, ibid., vol. 5, pt. 18, Apr., 1908, p. 104 (presumed poisonous bite). Deraniyagala,
Ceylon Journ. Sci., sec. B, vol. 16, 1931, p. 167.

Mabuya carinata Smith, The Fauna of British India including Ceylon and Burma ; Reptilia
and Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp. 206-207, fig. 68.

Tiliqua rufescens (part) Giinther, The Reptiles of British India, 1864, pp. 79-80 (Ceylon).

This large species is represented in the collections by U. S. N. M.
No. 120325 from Clodagh Estate, Rattota, Matale District, col-
lected by H. G. Deignan; and EHT-HMS Nos. 30504-30505, 30489,
30493 collected 12 mi. N of Trincomalee, Ceylon, by E. H. Taylor.
The specimens are typical. This species has a range extending
throughout Ceylon, and it is widespread in India and Burma.

The species may be recognized by the following characters:
supranasals separate; frontonasal longer than broad; prefrontals
separate (elsewhere they may be in contact) ; a pair of nuchals; no
postnasal; anterior loreal higher, but shorter, than second (the two
fused in No. 30490) ; three or four large scales on lower eyelid; post-
mental usually touches a single median scale, thus separating the
first pair of chinshields; dorsal scales subequal, 30 scale rows
around body (30-34 reported elsewhere); the temporals keeled;
scales keeled with three (young), five or, rarely, seven keels on
dorsum and side of adults; ventrals smooth; 15 to 18 smooth lamellae
under fourth toe. Reaches a length of 125 mm. snout to vent.
Olive above, uniform or with flecks or lines on the scale edges. The
dorsolateral light stripe from eye to tail often orange or reddish in
males during the breeding season.

* It is possible that the variation is not fully evident in young specimens as suggested under
Mabuya macularia Blyth (this paper).

490 The University Science Bulletin

Mabuya madardszi Mehely

Mabitia madardszi Mehely, Termes. Fuzetek, vol. 20, 1897, pp. 59, 61 (type locality, Kala-
Wewa and Madatugama, Ceylon).

Mabuia macularia (part) Smith, The Fauna of British India including Ceylon and Burma,
Reptilia and Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp. 264-266.

The distinctive modification of the squamation on the posterior
part of the thigh of this form caused me to believe at first that the
series here reported constituted a new form. However since the
specimens agree with the type description of Mabuya madardszi in
other characters, I suspect that the thigh characters were overlooked
by Mehely, as well as others who have seen Mehely's types, or
other specimens of this species from Ceylon.

The larger part of the specimens have heavy infestations of a
small mite on the back of the thigh that tends to distort the appear-
ance of the area. However when the mites are removed the char-
acters are distinct. Some of the specimens have no such infesta-
tions, and the scales are modified in the same manner.

I append a rather detailed description of a specimen to supple-
ment the original type description.

Diagnosis. A medium-sized, arboreal Mabuya, five- to seven-
keeled ; transparent part of eyelid broken up into at least five quad-
rangular scales; supranasals separated; prefrontals and parietals
also separated; five superciharies; no postnasals; two or three pri-
mary temporals; subcaudals not widened; chinshields broadly in
contact; an irregular series of scales covering a somewhat swollen
area on posterior thigh surface; posterior to insertion of leg, an
area of smaller irregular scales, and in the distal end of femoral
region another area of smaller scales, the two areas connected by a
double or triple series of large, somewhat irregular loosely imbri-
cating scales.

Description of EHT-HMS No. 30566. Portion of rostral visible
from above nearly two thirds of length of frontonasal, the two scales
forming a suture that separates the narrow elongate intemasals;
frontonasal distinctly wider than long, touching anterior loreals lat-
erally and frontal posteriorly, thus separating prefrontals; frontal
elongate touching only second supraocular, which is also in contact
with prefrontal; interparietals a little more than half length of
frontal; interparietal definitely longer than wide; nuchals and part
of parietals destroyed; nasal small, the part behind nostril greatly
reduced; anterior loreal much higher but less than half as wide as
posterior loreal; two presuboculars ; height of subocular in its length,
21/2 times; four supralabials anterior to subocular; four supraoculars;

Taylor: Ceylonese Lizards 491

five superciliaries ; four postsuboculars diminishing in size; two
primary temporals, three secondary temporals; mental almost equal
in length to postmental; first pair of chinshields in contact, second
pair separated by one scale, third pair by parts of three scales;
eight infralabials; three or four small ear lobules; diameter of cir-
cular ear opening half length of subocular; lower eyelid with a semi-
transparent area covered by five or more vertically elongated quad-
rangular scales.

Thirty scale rows around middle of body, the twenty dorsal and
lateral rows are seven-keeled (sometimes five in young) ; a varied
number (two to five) of keels on upper side of arm and leg; usually
three or more keels present on basal third of tail; scales following
nuchals somewhat irregular.

Limbs long, the adpressed hind limb reaching to axilla; lamellae
under fingers and toes not or bluntly keeled; 15-15 lamellae under
fourth toe; two median preanals distinctly larger than surrounding
scales, the three lateral preanals diminishing in size; subcaudals
not distinctly widened. Scales on back of thigh strongly modified;
a group of small scales behind insertion and another group of smaller
scales at distal end of femoral region; the scales between these areas
soft, more or less pointed, loosely imbricated, often nearly erect.

Color in life. Upper surface of head coppery brown, the eight dor-
sal scale rows copper colored with some olive wash and metallic re-
flections, and with some trace of darker markings in the middle of
each scale suggesting indistinct darker lines; an indistinct dorso-
lateral light line continues some distance on tail; a broad dark
brown band begins behind eye and continues far on tail, laterally
occupying two whole rows and parts of two other scale rows. A
cream line beginning on upper labials passes below ear, bordering
the brown band below and continuing some distance on the tail;
ventral surface washed lightly with dull olive.

Measurements in mm. Snout to vent, 72; tail, 99, partly regen-
erated; width of head at ear, 11.2; length of head to end of inter-
parietal, 12.5; snout to ear, 15; snout to arm insertion, 27.5; axilla
to groin, 35.5; arm, 21.5; leg, 35.2.

Variation. As regards the diagnostic scale characters, there is
little or no significant variation. The coloration, however, does vary.

No. 30570. This specimen has eight dorsal scale rows and the top
of the head is bright copper with metallic reflections. The borders
of the brown band above and below were greenish cream in life.
Nos. 30505 and 30571 have this identical coloration although they

492 The University Science Bulletin

differ much in size (respectively 72, 63, and 51 mm. in snout-to-vent
length). One of the specimens (30568) has the median dark areas
in the scales somewhat intensified and there are five narrow dark
dorsal lines evident when the specimen is submerged. Scale rows
vary between 30 and 32; only one has the latter count, and three
have 31. The others have 30 rows.

Two adult female specimens (30569, 30572) differ in having the
scales in the nuchal and temporal regions dimly keeled or smooth,
and the specialized scales on the posterior part of the thigh are more
definitely pointed and soft; the hind legs fail to reach quite to the
axilla. One female is strongly infested with small mites in the post-
femoral region.

The lamellae under the fourth toe vary between 15 and 18, 16
being most frequent. The number 18 was encountered once, 15 three
times. In none of the specimens is there a trace of ocellated spots
on the sides of the neck and body.

Remarks. It seems probable that this species has been confused
with another or other species of which macularia is the most prob-
able. The latter species has been reported and certainly occurs in
Ceylon. A Ceylon specimen is fortunately at hand and a description
is given.

Genus Riopa Gray

Riopa Gray, Ann. Mag. Nat. Hist., vol. 2, Jan., 1939, p. 332.

Genotype. Lygosoma punctatum.

Two species occur on Ceylon: Riopa punctata and a new form
here described, Riopa sing ha.

Key to the Species of Riop.a in Ceylon

Scales 24-26 rows, each dorsal and lateial scale with a dark spot. In young spots
forming 6 dark lines separated by light lines ; tail uniform red ; dorsolateral light
lines from rostral Riopa punctata

Scales in 28 rows; four very narrow dark lines on median scale rows; dorsolateral lines
from supraoculars ; vertical rows of white spots on neck and scattered white spots
on the sides ; a dorsolateral line from nuchal Riopa singha

Riopa punctata (Linne)

Fig. 1

fLacerta punctata Linne, Syst. Nat., 1, p. 369.

Scincus punctatus Gmelin, Hist. Amphib., 1799, p. 197, based on Seba's fig. 2, pi. 12, fig. 6.
Riopa punctata Smith, Fauna of British India including Ceylon and Burma, Reptilia and
Amphibia, vol. 2, Sauria, 1935, pp. 318-319.

U. S. N. M. No. 29412, "Ceylon"; EHT-HMS Nos. 30174-30181,
12 miles north of Trincomalee, Ceylon; E. H. Taylor, coll.

The series from near Trincomalee on the northwest coast, for the
most part, was found in coconut groves near the seashore. Indi-

Taylor: Ceylonese Lizards

493

viduals were secretive and their presence was usually made known
by the flash of the bright red tail. This red is present on the tails
of all young and half-grown specimens, but tends to become com-
pletely obsolete in adult animals.

A B

Fig. 1. Riova punctata (Linne). EHT-HMS No. 30178, 12 mi. N Trincomalee,
Ceylon. A. Head, dorsal view. B. Head, lateral view. X 6.

The color markings are strongly delineated. Two dorsolateral
cream lines begin on the tip of the snout and continue to the tail
where they become lost, the intervening dorsal ground color being
light olive-tan with six longitudinal series of black spots, one spot
on each scale. Seven lateral rows of scales likewise bear black
spots. The three upper rows are composed of larger spots and to-
gether suggest a dark lateral band or stripe. The ventral scale
rows, chin and throat are immaculate. In the young the tail, save
in the basal region, is unspotted and of a uniform shade of red.
As they grow older, the caudal scales develop small spots (even
those on the ventral surface) and gradually the red (sometimes
pink) coloration disappears. In the young the black spots are con-
tiguous forming continuous lines. On the head the dots are carried
forward to the snout but later tend to form a more or less sym-
metrical pattern on the head scales.

Scale counts about midbody are 24 to 26; however, when the
count is made a little farther forward sometimes 28 may be found.
The reason for the variation is that the axillary rows (32 about

494

The University Science Bulletin

body in axilla) tend to drop out quickly but one or two rows on
each side may continue back to near midbody. The scales in a
row from the nuchal to a point above anus number 68 to 76; the
scales under the tail to tip, number 90-91 (2 counts). In all, the
prefrontals are separated and the internasals are in contact (barely
contiguous in one specimen) .

U. S. N. M. No. 29412 is an old faded specimen agreeing with the
described series in all pertinent details.

In the figured specimen a small postnasal is evident. This occurs
in none of the other specimens, and is absent on the left side in the
specimen figured. I regard its presence as anomalous.

Riopa singha, sp. nov.

Fig. 2

Type. U. S. N. M. No. 29411, "Ceylon"; D. L. Karcher, coll.

Diagnosis. A species related to Riopa albopunctatum; four very
narrow dorsal lines on middle of back; wide dorsolateral cream lines
begin at supraoculars; below dorsolateral light line an irregular
dark stripe not of solid color, widest at neck and above arm and
axillary region tending to form four, fine, broken, dark lines pos-
teriorly. On this dark stripe vertical series of white dots alternating
with darker spots, these often somewhat irregular. Scale rows
around body, 28; a transparent eye disk present; 70 scales on back
between nuchals and a point above vent; snout to arm insertion
length contained in axilla to groin length 2% times; adpressed
limbs separated by a distance equal to one and a half to one and
two thirds length of arm.

A B

Fig. 2. Riopa singha sp. nov. U. S. Nat. Mus. No. 29411, Type.
A. Head, dorsal view. B. Head, lateral view. X 5.

Taylor: Ceylonese Lizards 495

Description of the type. Rostral separated from frontonasal by
paired siipranasals; frontonasal broader than long, forming suture
with frontal; prefrontals rather small, separated; frontal elongate,
longer than distance to snout but about two mm. shorter than parie-
tals, interparietal and frontoparietals together; parietals broadly in
contact behind interparietal, and bordered by a large temporal and
a pair of narrow nuchals; four 'supraoculars; eight superciliaries;
nostril in a small nasal; no postnasal; two loreals, anterior higher
than wide, higher but shorter than second which is nearly square;
seven upper labials, the suboculars not elongated; two primary
temporals, two or three secondaries, upper scales small; the lower
secondary separated from ear by four scale rows; seven lower
labials; mental followed by a large postmental and this followed
by three paired chinshields, first pair in contact, second pair sepa-
rated by one scale, third pair by five scales ; 28 scale rows about mid-
dle of body; about 36 rows around body at axilla; 72 scales from
parietal to above vent; limbs short, when adpressed, separated by
18 scales; 14 lamellae under fourth toe, strongly keeled; ear open-
ing large, about one third of eye opening; one large and one small
auricular lobule; eyelid with a more or less transparent disk; six
preanals all somewhat enlarged; tail regenerated; distance from
snout to insertion of arm contained in axilla to groin distance 2i/^
times.

Color. The specimen is of a dirty olive color and probably shows
none of the original coloration. A dorsolateral light line two and
one half scales wide begins on supraoculars and passes back onto tail;
the four median dorsal scale rows with hair-fine brownish lines
along their middle; a broad lateral black or brown stripe, tending
to form three fine lines posteriorly, and anteriorly with round light
spots sometimes forming vertical rows on sides of neck; below uni-
formly unspotted; limbs with brown spots above.

Measurements in mm. Snout to vent, 44; snout to forelimb, 11.4;
axilla to groin, 29; arm, 6.5; leg, 11.

Remarks. The type is in a good state of preservation save that the
abdomen has been opened widely, and one limb broken. It is a fe-
male as evidenced by several ovarian eggs. The tail shows at least
two different regenerations. There is a possibility that this form is
the representative of Riopa albopunctatum on Ceylon. The points
that appear to separate the species are the different body propor-
tion (snout to arm contained in axilla to groin 2% times, and the
limbs separated by a distance equal to one and one half times the

496 The University Science Bulletin

length of the arm) and some differences in color and markings (fine
dark lines on four medial scale rows, the wide dorsolateral light
line one and one half scale rows wide) .

The actual extent of differentiation between the two forms can be
determined only with more material from Ceylon and a study of the
Indian specimens of R. albopunctata with a view of determining
geographical trends in squamation'.

Genus Dasia Gray

Dasia Gray, Ann. Mag. Nat. Hist., vol. 2, 1839, p. 331.

Genotype. Dasia olivacea.

A single endemic species is known from Ceylon.

Dasia haliana (Haly and Nevill)

Euprepes halianus Haly and Nevill, Taprobanian, vol. 2, 1887, p. 56 (type locality Hena-
ratgoda and Anuradhapura, Ceylon); Boulenger, Fauna British India, including Ceylon and
Burma ; Reptilia and Batrachia, 1890, p. 213 ("probably belongs to Lygosoma") ; Haly, Ceylon
Admins. Rept., 1893, p. 13.

Theconyx halianus Annandale, Spolia Zeylanica, vol. 3, 1906, p. 191, figs. 1-4.

Lygosoma (Keneuxia) halianus Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 15, 1931, p.
174, pi. 37.

Dasia haliana Smith, Fauna British India, including Ceylon and Burma ; Reptilia and
Amphibia, vol. 2, Sauria, 1935, pp. 278-279.

This arboreal species is known from Central Province, North
Central Province, Southern Province, Western Province and North-
ern Province. It may readily be distinguished from other scincoid
species by the following characters: supranasals present; the nos-
tril in a single nasal; the prefrontals, interparietal, and frontopari-
etal distinct; the ear opening small; large, well-developed clinging
limbs; and a yellow-olive color with five or six broad transverse
black bands. No specimens are present in the collections.

Genus Sphenomorphus Fitzinger

Sphenomorphus Fitzinger, Systema Reptilium, 1843, p. 23.

Genotype. Lygosoma melanopogon.

Seven species from Ceylon are referred to this genus.

Key to the Species of Sphenomorphus in Ceylon

1. Frontoparietal divided 2

Frontoparietal single 6

2. Less than thirty scale rows about body 3

Forty scale rows about body dussuvneri

3. Parielals enclosing interparietal 4

Parietals separated by interparietal megalops

4. Dorsal scales striated deignani

Dorsal scales unstriated 5

5. Adpressed limbs barely overlap, prefrontals forming a common suture. . . taprobanense
Adpressed limbs separated by seven scales; prefrontals usually separated,

striatopunctatus

6. Males with side of head and throat blue-black, each scale with a whitish spot. . fallax
Males with throat bright rosy red, lacking black color and white spots rufogulus

Taylor: Ceylonese Lizards 497

Sphenomorphus dussumieri (Dumeril and Bibron)

Lygosoma dussumieri Dumeril and Bibron, firpetologie Generale . . . vol. 5, 1839, p.
725 (type locality, Malabar, India).

Lygosoma ( S-phenomorphus ) dussumieri Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 16,
1931, p. 169.

This species has been recorded by Deraniyagala from Peradeniya,
Ceylon. It also has a considerable range in southwestern India, and
in certain areas it is very common. It may be differentiated from
the other species of the genus by the following characters:

The distance between the snout and the forelimb is equal to, or a
little less than the axilla-to-groin distance; the prefrontals are
broadly in contact ; and four or five supraoculars are present. There
are 40 scale rows around the middle of the body, the dorsal scales
being striated; 20-25 lamellae are present under the fourth toe.

Sphenomorphus megalops (Annandale)

Lygosoma megalops Annandale, Spolia Zeylanica, vol. 3, 1906, p. 190 (type locality, Piit-
talam and Kitulgala, Ceylon); Smith, Fauna of British India, etc., Reptilia and Amphibia,
vol. 2, 1935, p. 289.

The following characters differentiate this species. The types
are said to be lost and no other specimens are known to be in col-
lections.

The length from the snout to the forelimb is contained one and
one half times in the axilla to groin length. The limbs overlap
when adpressed. The eye is large, its diameter as long as the snout.
There are no auricular lobules or denticulations on edge of ear.
There are no supranasals present, and the parietals are separated
by the interparietal. The dorsals and lateral scales are smooth but
the ventral scales are feebly keeled. These are arranged in twenty-
four to twenty-six scale rows about the body. The color is uniform
dark brown. The length of body of the type is two inches and of the
tail, two and three eighths inches.

Sphenomorphus deignani, sp. nov.

Fig. 3

Type. U. S. N. M. No. 120326, collected on Mount Ganoruwa
(Gangarowa?) Peradeniya, Kandy District, Central Province, Cey-
lon, by Herbert G. Deignan, 1944.

Diagnosis. A medium-sized, pentadactyl skink, characterized by
short limbs, the toes barely overlapping when adpressed ; ear-open-
ing large, the tympanum deeply sunk; paired frontoparietals, pre-
frontals broadly in contact; dorsal scales each with two striae;

31—90

498

The University Science Bulletin

nuchals lacking; 24 scales about body; numerous lateral scales with
white spots; jaws strongly barred with black.

Description of type. Rostral seen from above little more than
half width of frontonasal and forming a rather straight transverse
suture with latter; no supranasals; prefrontals large, broadly in
contact; frontal a little shorter than combined length of fronto-

B

Fig. 3. Sphenomorphiis deignani sp. nov. U. S. Nat. Mus. No. 120326, Type.
A. Head, dorsal view. B. Head, lateral view. X 5.

parietals and interparietal; four large supraoculars; frontoparietals
two; parietals inclosing interparietal; enlarged nuchals lacking; each
parietal bordered behind by an enlarged temporal; nine supercil-
iaries; nostril in a single nasal; two loreals of nearly equal size
and height; two preoculars, the upper small; suboculars eight or
ten, forming a well-defined row along the lower edge of eyelid ; eye-
lid covered with small scales ; diameter of eye equal to its distance
from nostril; a single primary temporal, not touching the parietal,
followed by a pair of secondary temporals, the lower of which is
separated from ear by three scale rows; seven upper labials; five
lower labials; the mental rather shortened but as wide as rostral,
one third as long as postmental; first pair of chinshields in contact;
second pair, separated by a scale, touching labials; third pair sepa-
rated from each other and the labials.

Taylor: Ceylonese Lizards 499

Ear-opening about one fourth of eye diameter, with one or two
minute thin lobules on its anterior border.

Twenty -nine scale rows around constricted part of neck; 34 about
body behind arm; 28 rows of subequal scales about middle of body;
median preanal scales somewhat enlarged; median subcaudals not
widened; 48 scales on back from parietals to point above vent.
Dorsal and lateral scales with distinct striations, three striae on
scales on nape, two striae on most of the other scales; ventral scales
smooth; striae present on ventral, lateral, and dorsal scales of tail,
but body ventrals have them barely indicated; greater part of tail
missing.

Limbs short; the arm reaches forward to eye; hind limb overlaps
the adpressed arm by half length of third finger; nineteen to twenty
lamellae under fourth toe.

Color and markings. Above dark olive brown, growing light
brown on back of body and base of tail; below on chin, abdomen,
and under side of tail creamy white ; a series of vertical black spots
or lines on jaws and on temporal region; sides of base of tail with
a few black flecks; heavy black reticulation with whitish spots on
arms and legs. Under a lens each scale of back and sides with a
crenellated, curved, black spot; together the spots form a complete
reticulation, especially evident when epidermis is shed. Many
lateral scales have minute, greenish white flecks.

Measurements in mm. Snout to vent, 55 ; tip of snout to ear, 12 ;
tip of snout to arm insertion, 20; axilla to groin, 28; arm length, 12;
leg length, 16.

Comments. The described species may be separated from the
other related Ceylonese and Indian species of Sphenomorphus as
follows: From megalops by the smooth instead of the keeled ven-
trals, seven instead of six upper labials, the presence of striae on
the scales, and the different coloration; from fallax and rufogulus
by the paired, rather than single, frontoparietal. From striato-
punctatus it differs in having the prefrontals broadly in contact
instead of widely separated, the limbs touching instead of being
separated considerably when limbs are adpressed, toes longer, the
lamellae under fourth toe nineteen to twenty instead of ten to
twelve, and the markings are different. From taprohanense, the
present form differs in coloration and markings, heavier and propor-
tionally longer limbs with more numerous scales on feet and palms,
and in the presence of striae on the scales. It is a much heavier
species. The type of taprohanense was said to have six lines of

500 The University Science Bulletin

black dots on the back, the sides of neck and body of a darker brown
minutely dotted with white.

The species is named for Dr. Herbert G. Deignan of the U. S.
National Museum, who collected the type of the species.

SphenomorphiLs taprobanense (Kelaart)

Eumeces taprobanense Kelaart, Prodromus Faunae Zeylanicae, vol. 2, pt. 1, 1854, p. 21
(type locality, Newera Ellia [= Nuwara Eliya], Ceylon); Giinther, Reptiles of British India,
1864, p. 89 (part, but not figure).

Lygosoma taprobanense Boulenger, Catalogue of the Lizards of the British Museum, vol. 3,
1887, p. 319; and Fauna British India, Reptilia and Amphibia, 1890, p. 206; Smith, Fauna
British India . . ., Reptilia and Amphibia, vol. 2, Sauria, 1935, pp. 287-288.

Lygosoma ( Sphenoniorphus ) taprobanensis Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 16,
1931, p. 120.

This form, of which I have no specimens available, may be dis-
tinguished by the following characters: The distance between the
end of the snout and the foreleg is contained in axilla to groin dis-
tance one and one third to one and three fifths times. The pre-
frontals are in contact or separated. There are two subequal loreals
present. The ear opening is about one half the size of the eye open-
ing. The body scales are smooth, arranged in 24-26 scale rows
about the middle. The limbs just meet or narrowly fail to meet
when adpressed. The color is brown above with six dark longitu-
dinal lines formed by dots. The upper half of the flank and neck
are dark brown, while the color of the lower margin is not well de-
fined. The sides of the neck are with or without white spots. The
ventral surface is whitish save that the throat of the adult male is
dark blue or purple. The snout-to-vent measurement is 58 mm.

The species is found in the mountainous regions of the central and
southern parts of Ceylon.

Sphenomorphus striatopunctatus (Ahl)

Lygosoma punctatolineatum (not of Boulenger, 1893) Boulenger, Spolia Zeylanica, vol. 4,
1907, p. 173 (type locality, Hakgalla, Ceylon).

Lygosoma (Sphenomorphus) punctatolineolatus Deraniyagala, Ceylon Journ. Sci., sec. B,
vol. 16, 1931, p. 169.

Lygosoma striatopunctatum Ahl, Zool. Anz., vol. 6, 1925, 1 and 2, p. 20; Smith, Fauna
British India, including Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935, p. 288.

The species here described agrees with the form striatopunctatus
in practically all characters save that the prefrontals are moderately
large and in contact. The following description is from a Ceylon
specimen, No. 120327, in the U. S. National Museum, which, with
No. 120326, was collected by Dr. H. G. Deignan, Mount Ganoruwa,
Peradeniya, Kandy District, Ceylon.

Distance between end of snout and forelimb (15 mm.) contained
in axilla to groin distance (21 mm.) 1.4 times; snout moderately
long, oval; rostral convex, broadly in contact with frontonasal,

Taylor: Ceylonese Lizards 501

which is broader than long; prefrontals in contact (said to be rarely
separated) ; frontal as long as frontoparietals and interparietal to-
gether; parietals large forming a suture behind interparietal; two
frontoparietals; no nuchals; four large supraoculars, the last bor-
dered by two small scales; second largest, first and second touch
the frontal; eight or nine superciliaries; a large primary temporal
followed by two large secondary temporals, these bordered behind
by three smaller scales; seven upper labials, the fifth below eye;
presuboculars, suboculars and postsuboculars forming a continuous
series of eight or nine scales, all dark colored; two loreals, second
largest; nasal single; no postnasal; mental followed by an undi-
vided postmental and two pairs of large chinshields that border
labials but only first pair in contact; third pair rather large, sepa-
rated from labials by an elongate scale; diameter of ear opening
about one third that of eye, with one or two projecting lobules;
tympanum deeply sunk; body scales smooth; 26 scale rows about
middle of body; 53 scales from parietals to above vent; tail some-
what thickened at the base, definitely quadrangular in cross-sec-
tion; tail partly regenerated, the ventral scales being enlarged on
regenerated part (normal scales small) ; limbs when adpressed
separated by about seven scales ; thirteen lamellae under the fourth
toe; palms and soles with flattened, rather than conical, scales.

The color is as follows (male) : Entire head bluish black without
white spots, gradually becoming brown on neck; brown on body,
with indicated lines (many dorsal scales missing) ; slightly darker
on upper sides, many of the dark scales having a light spot or fleck;
lower scale rows on sides dirty white, as are the ventral abdominal
scales; tail colored like body, unspotted below.

Snout to vent, 40 mm.; arm, 7 mm.; leg, 11 mm.; axilla to groin,
21 mm.; snout to foreleg, 15 mm.; width of head, 5.6 mm.; head
length to back of parietal, 8 mm.

Sphenomorphus fallax Peters

Fig. 4

Lygosoma fallax Peters, Mon. Berlin Akad., 1860, p. 184 (type loealit.v, "Ratnapura, Trin-
comali" Ceylon); Boulenger, Catalogue of the Lizards of the British Museum (Natural History),
vol. 3, 1887, p. 320 (part); Fauna of British India, Reptilia and Batrachia, 1890, p. 206;
Mehely, Termes. Fuzetek, vol. 20, 1897, p. 61 (Kala Wewa, Ceylon); Smith, Fauna of British
India including Ceylon and Burma, Reptilia and Amphibia, vol. 2, Sauria, 1935, pp. 288-289.

Lygosoma (Sphrjwmorphut,) fallax Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 16, 1931,
p. 172.

Eumeces taprobancnsis (part) Giinther, Reptiles of British India, 1864, p. 89 (not figure).

The following specimens are in the collections: U. S. N. M. Nos.
27286, 29413, "Ceylon," and EHT-HMS Nos. 13082-13093, 23 miles
west of Trincomalee, Ceylon.

502

The University Science Bulletin

This small species is easily distinguished from Sphenomorphus
striatopunctatus, S. taprobanense, S. megalops, S. dussumieri and
S. deignani by having the frontoparietals fused together to form a
single scale. It is not entirely easy to distinguish from S. rujogulus,
a species described herein, that likewise has the frontoparietal
single. The males of fallax may be distinguished easily since fallax
has a blue-black head, and cream-white dots on the side of head
and throat; and rujogulus has a large red area on the throat and
no trace of the blue-black coloring on the head. However, females
lack these characters and the slight scale differences and the ab-
sence of well-defined color patterns make them difficult to distin-
guish. Twelve miles north of Trincomalee, rujogulus was found

A B

Fig. 4. Sphenomor-phus fallax (Peters), EHT-HMS No. 13084, 20 mi. W
Trincomalee, Ceylon. Male. A. Head, dorsal view. B. Head, lateral view.
X5.

alone and there it was a very common species in the forest near the
seashore. Twenty miles from Trincomalee to the west it was found
in the forest with fallax, the latter apparently in the greater num-
bers. A very large number of specimens of both species might very
readily have been taken had I so desired, as they seemed to be
everywhere on the forest floor.

Eight specimens of *S. fallax have the prefrontals forming sutures
of varying width, seven have them separated by variable distances.
None have true nuchals, the parietals being bordered by a temporal
and five or six body scales. In several specimens the two outer body
scales fuse to make a larger scale following the temporal, and re-

Taylor: Ceylonese Lizards 503

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504 The University Science Bulletin

ducing the number of scales to three or four. The frontal is usually
rather small (2.5 mm. in length) while the combined frontoparietal,
interparietal and the parietal length measures 4.15 mm. on the me-
dian line. The scales of the third pair of chinshields are separated
from each other by a single scale and from the labials by a single
scale. The presubocular, subocular and postsubocular series are
continuous, large, and heavily pigmented like the lateral head scales.

The coloration in the males is deep, almost uniform brown on
the ten median dorsal scale rows, with little or no trace of pattern
visible under the lens; no dorsolateral lighter line is visible. The
three lateral scale rows each show a very slight suggestion of three
light lines, often scarcely traceable. The underside of the body is
immaculate. The sides of the head, and throat, are bluish black,
the color extending to behind the ear opening, each scale with a
whitish or bluish white spot more or less clearly defined. The top
of the head is variable, dark or lighter brown. The blue-black
coloration is missing on head and throat in the females but the
lateral whitish lines from axilla to groin are rather clearly distin-
guishable, the upper being most distinct. A distinct dorsolateral
line two half scales wide is present, below which there is a distinct
dark band, one whole and two half scale rows wide. Some of the
cheek scales and labials may show lighter areas. The sides and
ventral part of the tail are more or less heavily flecked with dark
brown.

Further data on this series of specimens is given in the following
table. Measurements indicate a relatively small amount of size
variation other than that dependent on sexual variation. This is
evident in the wider head in the males and the somewhat slenderer
bodies in the females. None of the males have complete tails.

Sphenonwrphus rufogulus sp. nov.

Fig. 5

Type. EHT-HMS No. 30229 collected 12 mi. N of Trincomalee,
Ceylon, Sept. 1944, E. H. Taylor, collector.

Paratypes. EHT-HMS Nos. 30195-30228, 12 mi. north of Trin-
comalee, Ceylon, Sept., 1944; Nos. 30230-30233, 21 mi. east of
Trincomalee, Ceylon, Sept., 1944, E. H. Taylor coll.

Diagnosis. A small species of the size of Sphenomorphus jallax,
and, like it, having the frontoparietals fused, the interparietal and
the internasals enclosed by the parietal, two or three primary
temporals, but differing in having the throat and chin lacking dark

Taylor: Ceylonese Lizards

505

pigment, and colored bright red in the males. Males with three to
five dim ocelli on the last supralabials and temporals. Dorsal color-
ation lighter brown.

Description of the type. Rostral broad, low, two and one half
times as wide as high; frontonasal one and one half times as broad
as long; prefrontals large, forming a broad sutm^e; frontal longer
than its distance from end of snout, a little shorter than frontoparie-
tal; width of frontal contained in width of supraoculars one and
one fourth times; frontoparietal a single scale, its length minutely
less than its width; interparietal small, enclosed by large parietals;

A B

Fig. 5. Sphenomorphus rufogulus sp. nov. EHT-HMS No. 30228, Paratype
and topotype. A. Head, dorsal view. B. Head, lateral view.

combined length of parietals and frontoparietal exceeds their dis-
tance from the snout tip about one eighth; nostril in a single nasal
followed by an anterior loreal wider at top, higher but narrower than
posterior loreal; a pair of preoculars, the lower larger; seven or eight
superciliaries ; four supraoculars, two touching frontal; seven supra-
labials, the fifth below middle of eye; a series of six scales lie be-
tween labials and small scales of eyelids; a rather large scale
partly between fourth supraocular and parietal; one rather large
postocular; three primary temporals, the upper largest; two sec-
ondary temporals, the lower separated from ear by two scales ; men-
tal more than half as wide as postmental ; three pairs of chinshields,
first pair in contact, second separated by a scale but touching

506 The University Science Bulletin

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Taylor: Ceylonese Lizards 507

labials; third pair separated by a scale and each separated by a
single scale from labials; five elongate infralabials.

Scales smooth, in 28 rows around middle of the body; median
caudals not enlarged; 48 scales from parietals to above anus; 40
scales from axilla to groin; 16 lamellae under fourth toe with an
indistinct keel or tubercle on each; adpressed hind limb separated
from adpressed arm by four or five scales.

Color. Above, light wood-brown, the head not or very slightly
darker than back. The dorsolateral line (present in females) is
barely indicated on sides of body and neck; on tail represented by
black dots on two scale rows ; temporal region with some flecks and
several small ocellated cream spots on temporals and posterior la-
bials ; a bright red area on chin and neck

Measurements in mm. Snout to vent, 41 ; tail, 41 (tip missing) ;
arm, 8.2; leg, 12.3; axilla to groin, 22.2; snout to arm insertion, 15.3;
head width, 6.1; head length, 8.

Remarks. It is entirely probable that this species is identical
with the form figured by Giinther in his Reptiles of British India,
1864, pi. 13, fig. B, as Eumeces taprobanensis. The specimen shows
two primary temporals, a rather common condition in rujogulus and
it is presumed that the frontoparietals are fused. At any rate the
figure might serve very well as an illustration of this species, es-
pecially for the females. Boulenger (Catalogue, vol. 3, pp. 319-320)
however has referred the figure to the true taprobanense and some
of the other specimens mentioned by Giinther to fallax.

A table of measurements and scale counts of a part of the para-
type series is given.

Genus Chalcidoseps Boulenger

Chalcidoseps Boulenger, Catalogue of the Lizards in the British Museum, 2d ed., vol. 3,
1887, p. 423.

Genotype. Chalcidoseps thwaitesi.

This genus has the nostril pierced in the rostral near the posterior
edge. A scale lies above the first labial preceding the loreal ; a very
large frontonasal is present preceded by a relatively small rostral;
no prefrontals or frontoparietals present. The body is elongate,
with four small limbs each bearing four digits

508 The University Science Bulletin

Chalcidoseps thwaitesi (Giinther)

Nessia thwaitesi Giinther, Ann. Mag. Nat. Hist., ser. 4, vol. 9, 1872, p. 86 (type locality,
Ceylon).

Chalcidoseps thwaitesi Boulenger, Catalogue of the Lizards of the British Museum, 2d ed.,
vol. 3, 1887, p. 423, pi. 38, fig. 1; and Fauna of British India, including Ceylon and Burma;
Reptilia and Batrachia, 1890, p. 226, figs. ; Dereniyagala, Ceylon Journ. Sci., sec. B, vol 16, (2),
1931, p. 176; Smith, Fauna of British India, etc., Reptilia and Amphibia, vol. 2, Sauria, 1935,
pp. 335-336, fig. S3.

The genus has but a single species, which may be distinguished
from any other known Ceylonese lizard by the generic characters
listed above. It is known only from Ceylon specimens having been
taken at Gammaduwa, in the Central Province. It is said to live
at from 4,000 to 5,200 feet elevation among dead leaves and other
vegetation.

Genus Nessia Gray

Nessia Gray, Ann. Mag. Nat. Hist., vol 2, 1839, p. 336.

Evesia Gray, Ann. and Mag. Nat Hist., vol. 2, 1839, p. 336. Genotype. Evesia mono-
dactylus Gray (type locality unknown; presumably Ceylon.)

Pseudodactylus Fitzinger, Systema Reptilium, 1843, p. 23. Genotype. Evesia bellii
Dumeril and Bibron (= N. monodactyla).

Tetrapedos Jan, Arch, fiir Naturg., Berlin, 1860, p. 69. Genotype. Tetrapedos smithii
Jan. from Ceylon.

Acontias (part) Boulenger, Catalogue of the Lizards of the British Museum, vol. 3, 1887,
p. 224.

Anguiniccphalus Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 18, 1934, p. 232. Genotype.
Acontias layardi Kelaart, from Ceylon.

Genotype. Nessia burtoni Gray (type locality unknown; pre-
sumably Ceylon).

The group of species considered under this genus are undergoing
a variety of evolutionary changes pointing to greater specialization.
Usually loss or reduction of limbs, loss of an ear-opening, the pres-
ence or absence of toes are regarded as generic characters and such
designations have been proposed. However, in the case of Nessia,
the conformity of the scale patterns of the various species, the great
similarity of the distinctive head scales seem to warrant the reten-
tion of a single genus for the series of varied forms. It is presumed
that the genus has undergone its entire evolution on the Island of
Ceylon since it is not known elsewhere. The closest Asiatic relative
seems to be Chalcidoseps.

The genus Brachymeles comprises a group of species comparable
to these forms, confined so far as is known to the Philippine Islands.
There the genus is represented by some ten known species. They
are widely distributed in this archipelago, being absent so far as is
known however, in the Calamianes Islands and Palawan. The
more generalized, more primitive forms, having typical, well-de-

Taylor: Ceylonese Lizards 509

veloped pentadactyl limbs are still extant. Brachymeles schaden-
bergi typifies this condition.*

One might account in part for the diversity of forms of Brachy-
meles by the archipelagic condition, obtaining at various times in
geological history of the Philippines, that has allowed the factor of
isolation effective play. No similar archipelagic condition exists
now and it is doubtful if it has ever existed so as to play an effec-
tive part in the evolution of Nessia in Ceylon. One must suppose
that the isolated elevated areas have provided "isolation" necessary
for their diversification.

Key to the Species of Nessia

1. Two or four limbs present; interparietal broader than frontal (except sarasi-

norum) ; ear opening present 2

Limbs absent 6

2. Limbs bearing clawed digits 3

Limbs budlike, lacking clawed digits 4

3. Four limbs present, tridactyl ; 24 scales at midbody burtonii

Four limbs present, didactyl; 24 scales at midbody didactyla

4. Four limbs present ; 24-26 scales about midbody monodactyla

Two limbs present ; scales variable 5

5. Scale rows about midbody, 28 hipes

Scale rows about jnidbody, 22 ; interparietal narrower than frontal sarasinorum

6. Snout flattened below, projecting sharklike; frontonasal one third width of ros-

tral ; ear-opening present hickanala

Snout not especially flat; not sharklike; frontonasal more than half length of
rostral 7

7. One large elongate loreal ; preoculars small ; frontonasal nearly as long as rostral,

layardi
Two loreals, the posterior lower than anterior; frontonasal a little more than half

of rostral length deraniyagalai

Nessia burtonii Gray

Nessia burtonii Gray, Ann. and Mag. Nat. Hist., vol. 2, 1839, p. 386; Kelaart, Prodromus
Faunae Zeylanicae, vol. 2, pt. 1, 1853, pp. 11-12; Gilnther, Reptiles of British India. 1864,
p. 97.

Acontias burtonii Boulenger, Catalogue of the Lizards of the British Museum, vol. 3, 1887,
p. 425 ; and Fauna British India including Ceylon and Burma, Reptilia and Amphibia, 1890,
p. 227; Smith, Fauna of British India . . ., Reptilia and Amphibia, vol. 2, Sauria, 1935,
pp. 357-358.

Acontias (Nessia) burtoni Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 16, (pt. 2), 1931,
p. 177; and idem, vol. 18, pt. 2, May 22, 1934, pp. 231-232.

Description. Snout subacuminate, at least twice as long as the
orbit, strongly projecting beyond the lower jaw; rostral about two
fifths as long as the snout; fronto-nasal longer than the rostral, dis-
tinctly narrower in front than behind ; frontal longer than the fronto-
nasal, mesially notched on each side by the first supraocular; in-
terparietal broader than the frontal; parietals narrow, obliquely
placed, in contact behind the interparietal bordered on each side by

* Taylor, in Lizards of the Philippines, 1922, p. 22, speaks of Brachymeles schadenbergi as
the most specialized. This is lapsus; "least specialized" is intended.

510 The University Science Bulletin

an elongated temporal shield; 4 supraoculars, the first two in contact
with the frontal; 5 superciliaries, the first much the largest; one
long loreal sometimes divided in the middle; a preocular; lower eyelid
an opaque disk or partly divided into scales ; upper eyelid vestigial ;
4 supralabials, the first very long, the second below the eye; mental
large with an azygous shield behind it; ear-opening punctiform.
Body very elongate; scales smooth, 26 to 28 round the forepart of
the body, 24 round the middle, dorsals largest; preanals not or but
feebly enlarged; limbs very short, tridactyle, clawed; forelimb origi-
nating at about the level of the 20th ventral scale, about as long
as four scales, shorter than the hind limb.

Light brown above, the scales tipped or edged with dark brown,
the general appearance being almost uniform brown or reddish-
brown; paler below.

From snout to vent 75 mm.; tail bluntly pointed, about three
quarters the length of the head and body.

Range. Central, Western and Sabaragamuwa Provinces." (from
Smith, loc cit.)

The species is reported by Kelaart from Ambegammoa and Ka-
duganava (several smaller specimens).

Nessia didactyla (Deraniyagala)

Acontias (Nessia) didactylus Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 18, 1934, pp.
232-233 (type locality, Polgehavala).

Nessia didactyla Smith, Fauna of British India including Ceylon and Burma, Reptilia and
Amphibia, vol. 2, Sauria, Feb. 7, 1935, p. 358.

Description. "Anguiniform, with didactyle, clawed limbs. Snout
bluntly acuminate, jaws overshot exposing anterior teeth of upper
jaw. Lower eyelid scaly, ear small, about 7 scales behind eye which
is midway between nostril and ear. Rostral covers about a third
of the snout, with the nostril pierced in its anterior region and con-
nected to the posterior margin of rostral by a groove. Frontonasal
longer than the rostral, slightly shorter than the frontal which is
shorter than the interparietal. The last is contiguous with the sec-
ond and third supraoculars. Parietals comparatively wide, wider
than the supraoculars and contiguous; behind them a row of 4 en-
larged nuchals in a single transverse series.

"Supraoculars 4, the first and second form emarginations in the
frontal. Supralabials 4, the first equals loreal and is 1.5 times
length of rostral, the second is subocular. Mental moderate, con-
tiguous with a single triangular chinshield behind which are three
pairs of elongate shields which are separated from each other mesi-
ally by a single row of scales. Infralabials three. Body scales di-

Taylor: Ceylonese Lizards 511

rected ventrally, 26 round mid body and about a hundred from
axilla to groin, 7 longitudinal pectoral rows. Anterior limb 23 scales
behind gape, as long as rostral with two well developed clawed digits
and seven scales along its edge exclusive of the digits, posterior limb
longer, with 10-13 rows of scales along its edge exclusive of the
digits. Preanals feebly enlarged. Caudals subequal. Extremity
of the cylindrical tail is feebly compressed dorsally.

"Colours. Brown, the dorsal scales outlined in darker, ventrally
lighter.

"Dimensions. Snout to ear 6.5 mm., gape 5 mm., ear to forelimb
7 mm., snout to cloaca 65 mm., tail 36 mm." (Type description.)

Range. Known only from the type locality, Polgehavala, Ceylon,
elev. 241 ft.

Nessia monodactyla (Gray)

Evesia monodactylus Gray, Ann. and Mag. Nat. Hist., vol. 2, 1839, p. 33G (type locality
unknown).

Nessia monodactyla Glinther, Reptiles of British India, 1864, p. 97.

Acontias monodactylus Boulenger, Catalogue of the Lizards in the British Museum, vol. 3,
1887, p. 425; Fauna of British India including Ceylon and Burma, Reptilia and Amphibia,
1890, p. 228.

Evesia bellii Dumeril and Bibron, firpetologie Generate, vol. 5, 1839, p. 782. (Substitute
name for monodactyla; same type.)

Tetrapedos smithii Jan, Arch. f. Naturg. Berlin, 1860, p. 69, pi. 2, figs. 4-12 (type locality,
Ceylon).

Acontias (Nessia) monodactylus Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 16, 1931,
p. 178.

Nessia monodactyla Smith, Fauna British India including Ceylon and Burma, Reptilia and
Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp. 358-359.

Three specimens are in the EHT-HMS Collection, Nos. 30057,
30058, 30059. The following are scale counts and measurements:

30056 30057 30058

Length of head to nuchals 6.3 6.1 5.8

Width of head 5.0 5.0 4.7

Width of body 5.5 5.7 4.8

Tail 46 29.8

Length to foreleg 15 14.2 13.2

Axilla to groin 65 66 48

Scales around neck 24-26 26 27

Scales on middle of body 26 24 24

Scales 1 cm. before anus 24 24 24

Ventral scales postmental to the anal scales 118 119 121

Subcaudals 85 83

Length of head and body 83 84 62

Length to ear opening 6.7 7 6.2

Conformation of the head scales agrees in general with the type
description. The temporal bordering the parietal is divided save
on one side in No. 30056 where it is single. In this same specimen

512 The University Science Bulletin

the parietals do not inclose the interparietal (an obvious anomaly).
The pineal eye is strongly evident in No. 30057. A groove or a pit
is present at the eyespot.

The anterior part of frontal is shorter in 30057 than in the other
two specimens. Six preanals are present. There are about nine
scales in a row on outer surface of limb. The loreal is larger than
the first labial, and is divided in none. Five supraoculars; five su-
pralabials ; four infralabials are present. The area posterior to the
anus is swollen, and covered with small scales.

As to the condition of the budlike limbs, those of Nos. 30056
and 30057 are shorter and broader than those in 30058. In the lat-
ter, the smallest specimen, the legs are somewhat longer, and taper
more toward the tips. This last specimen is from Peradeniya,
Ceylon. The other two are from Ceylon, but definite localities are
lacking.

Nessia bvpes Smith

Nessia (Evesia) smithi Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 18, 1934, p. 232 (type
locality, Gammaduva, Central Province, Ceylon).

A^essia bipes nov. nom. for A'essia (Evesia) smithi preoccupied, Smith, Fauna British India,
including Ceylon and Burma; Reptilia and Amphibia, Vol. 2, Sauria, Feb., 1935, p. 359.

This species has not been adequately described. It is said to agree
with monodactyla in the squamation of the head. It differs from
that species in having a budlike pair of posterior limbs only and in
having 28 scales around the body both anteriorly and in the middle.

From snout to vent 80 mm. Known only from the type specimen,
from Gammaduva, Central Province.

Nessia layardi (Kelaart)

Fig. 6

Acontias layardi Kelaart, Prodromus Faunae Zeylanicae, vol. 2, 1853, pp. 12-13 (type lo-
cality, "soil of Cinnamon Gardens of Colombo, Ceylon); and Ann. and Mag. Nat. History, (2),
13, 1854, p. 26 ; Boulenger, Catalogue of the Lizards in the British Museum Natural History,
2d ed., vol. 3, 1887, p. 426, and Fauna British India including Ceylon and Burma; Reptilia and
Amphibia, 1890, p. 228, fig. p. 227; Gunther, Reptiles of British India, 1864, p. 96.

Acontias (Nessia) layardi Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 16, 1931, p. 179,
pi. XXXVIII.

Acontias ( Avguimcephalus) layardi Deraniyagala, Ceylon Journ. Sci., sec. B, vol. 18, 1934,
p. 231.

Nessia layardi Smith, Fauna of British India, including Ceylon and Burma . . ., Reptilia
and Amphibia, vol. 2, Sauria, Feb. 7, 1935, p. 359, fig. 4, p. 356.

Differs from burtonii in the following particulars: Frontonasal
broader and shorter than the frontal ; three supraoculars, the first
only in contact with the frontal notching its lateral margin; first
superciliary larger, entering supraorbital region; a pair of nuchals
often present; no ear-opening; 24-26 scales around the fore part of

Taylor: Ceylonese Lizards

513

B

Fig. 6. Nessia layardi (Kelaart). Figure from Boulenger, Fauna of British
India . . ., 1890, fig. 63, redrawn. Much enlarged.

the body, 22 or 24 round the middle; no limbs. On each side of the
vent, in a depression of the body and more or less hidden by scales,
a minute horny tubercle can be discovered with a good glass ; it rep-
resents what is left of the hind limb.

A single specimen of Nessia layardi, U.K.M.N.H. No. 24135, was
collected at DambuUa in the northern part of Central Province by
Dr. W. C. Osman Hill.

The specimen, when compared to Boulenger 's figure (Fauna of
British India . . ., 1890, pp. 356-358, fig. 84) shows no differ-
ences of significance. However the interparietal is slightly less an-
gular (more curved) than the figure and the first lower labial is
shorter.

There are 120 scale rows (transverse) and 24 (anteriorly) to 22
(middle and posteriorly) scale rows. The total length of the eye is
less than half the length of the loreal. The small nuchals are present ;
no ear opening. Three supraoculars are present.

There is a slight depression on each side of the body near the
vent, covered with small irregular scales marking the point where
the limb formerly stood.

The range of the form is Central Province and Western Province
although the type locality in the latter province has been questioned
by Deraniyagala {loc. cit).
32—90

514

The University Science Bulletin

Nessia sarasinorum (F. Miiller)

Fig. 7

Acontias sarasinorum F. Miiller, Verh. Nat. Gesel. Basel, vol. 8, 1889, p. 702, pi. X (type
locality, Inamalua, Ceylon); Boulenger, Fauna British India, including Ceylon and Burma;
Reptilia and Amphibia, London, 1890, p. 228.

Acontias (Nessia) Sarasinorum Deraniyagala, Ceylon Joum. Sci., sec. B, vol. 16, part 2,
1931, p. 178.

Nessia sarasinorinn Smith, Fauna British India, including Ceylon and Burma; Reptilia and
Amphibia, vol. 2, Sauria, Feb. 7, 1935, p. 360.

A specimen of this rare species, EHT-HMS No. 30003, was taken
from under a log, near a small forest stream 21 mi. west of Trin-
comalee, Ceylon. Its movements were rather slow as it crawled in
a bm-row in the sandy earth.

The color was dull lavender to grayish lavender in life. Preserved
it is lead color, the head, between rostral and nuchals being dark
lead. The tail is regenerated, with a ventral light spot, and a dim,
incomplete lighter ring on scales preceding the beginning of the re-
generated part. The dorsal scales when closely examined show
darker areas. There is no external trace of the front limbs, but the
scale irregularities show the point where limbs were present in an-
cestral forms. The hind limbs are budlike, covered by four rows of
scales.

There are twelve scale rows about tail, 1 cm. behind anus; 24 rows
about neck; 22 about the middle of the body; 127 scales in a row
from postmental to anal; 124 scales on back from parietals to level
of hind limbs.

B

Fig. 7. Nessia sarasinorum (Miiller). EHT-HMS No. 30003, 21 mi. w Trinco-
malee, Ceylon. A. Head, dorsal view. B. Head, lateral view. About X 10.

Taylor: Ceylonese Lizards 515

The details of the head squamation follow: Snout extending be-
yond mouth, covered by a thickened shield much broader than long;
nostril pierced in the anterior lateral part with a suture extending
back to anterior loreal ; frontonasal much broader than long, broader
but shorter than the frontal; latter broadly hourglass-shaped; inter-
parietal small, triangular, narrower and shorter than frontal ; parietals
widened anteriorly, enclosing the interparietal ; each interparietal bor-
dered by two elongate scales, the medial of which may be regarded as
nuchals; two large loreal scales of equal height; four supraoculars;
a large postocular, one preocular; four superciliaries, very irregular
in size; four or three suboculars (the first may be excluded from
orbit). Scales of eyelids more or less fused together; four suprala-
bials, the first equal in size to the combined loreals; four infralabials;
mental thickened, followed by an azygous postmental, wider than
the mental; three well-defined pairs of chinshields, the two anterior
separated by a single scale ; ear opening present, minute.

Snout to vent, 86 mm.; tail (regenerated), 37 mm.; diameter of
head, 5 mm.; diameter of body, 5.5 mm.; snout to ear, 7 mm.

Smith {loc. cit.) calls attention to errors in Miiller's description.

Nessia hickanala Deraniyagala

Nessia hickanala Deraniyagala, Proc. Linnean Soc. of London, 1939-40, pt. 1, Feb. 9, 1940,
pp. 37-39, figs. a-d.

Description. Habit anguiniform, apodal, with a few small scales
indicating the position of the hind limbs. Snout very shark-like,
being convex dorsally, but strongly flattened ventrally with a
prominent labial ridge; jaws overshot, but front teeth not exposed.
Eye small, its lower lid covered by three transparent scales; ear
small, about six or seven scales behind eye which is slightly closer
to nostril than to ear. Rostral covers about three fifths of the
snout, with the nostril pierced in its anterior region and connected to
the posterior margin of this shield by a groove.

Frontonasal only about a third of the length of the rostral, and
about half the length of the frontal, which is shorter and narrower
than the interparietal. Parietals as wide as supraoculars and con-
tiguous behind the interparietal; behind them a single transverse
row of four enlarged occipitals. Supraoculars three, one large post-
ocular. Preocular strongly enlarged, cuneiform. Supralabials four,
the second is below orbit; mental moderate, contiguous with a
single triangular chinshield, and with four enlarged shields along
each mandible below the four infralabials, of which the last is con-
spicuously elongated. Body-scales directed ventrally; dorsals fee-

516

The University Science Bulletin

bly enlarged, 20-21 scales round midbody. Preanals conspicuously
enlarged; caudals subequal.

Color. Pinkish-brown, darker dorsally, each scale-pocket a dark
brown which becomes reduced with age. After death the pink is
replaced by a pale grey.

Measurements (of largest specimen). Snout to vent, 92 mm.;
around body in middle, 17 mm. (Of type: Snout to vent, 68 mm.;
snout to ear, 7 mm.) The tails of all known specimens were either
mutilated or regenerated.

The above description is taken almost wholly from the type de-
scription.

Nessia deraniyagalai, sp. nov.

Fig. 8

Type. EHT-HMS No. 30059 ^ , collected 16 mi. N Trincomalee
near shore, on small hillock at an elevation of about ten meters,
Oct. 24, 1944, E. H. Taylor, collector.

Diagnosis. A legless Nessia, differing from the legless A^. layardi
in having two, instead of one loreal, the anterior much the higher;
a frontoparietal much broader than frontal but only five eighths as
long; part of frontal anterior to lateral notch of much larger area
than part posterior to notch; frontoparietal wider than frontal;
second supralabial not higher than the much enlarged first. Scale

B

Fig. 8. Nessia deraniyagalai sp. nov. EHT-HMS No. 30059. 16 mi. N Trinco-
malee, Ceylon. Type. A. Head, dorsal view. B. Head, lateral view. X 10.

Taylor: Ceylonese Lizards 517

rows about body: anteriorly, 22, not extending to midbody, where
there are 20; 18, one cm. in front of tail.

Description of species. Snout covered with a much thickened ros-
tral plate, more than one and one half times as wide as long; nostril
pierced laterally, with a long suture from it passing back to the first
loreal; posterior lateral border of rostral somewhat emarginate;
frontonasal as wide as rostral, but somewhat shorter; wider than
frontal, but latter one and one half times as long; frontal narrowed
medially with short lateral sutures, the anterior part of greater
width and area than posterior part; frontoparietal wider than fron-
tal, angular posteriorly, convex anteriorly; parietals narrowed some-
what anteriorly, in contact posteriorly, bordered behind by a pair
of nuchals and an elongate temporal; two loreals, anterior higher;
five supraoculars (the anterior of which may actually represent a
greatly enlarged superciliary), (?) three or four superciliaries ; five
supralabials, first large, elongate, second borders orbit, separated
from eye by two minute scale rows on lower eyelid; a preocular; a
small presubocular and a postsubocular, the latter above third labial ;
four infralabials, the posterior elongated; mental thickened, extend-
ing back almost as far as the rostral seen from below, and distinctly
longer than portion of rostral extending beyond the mouth; post-
mental narrower and shorter, the posterior edge rounded somewhat;
three pairs of chinshields, the second widest, the two anterior sep-
arated by a single median scale. Lower eyelid movable, the eye
slit two thirds millimeter in length; no ear opening (however in
the normal ear position there is a slight lateral depression with one
or two minute scales) ; limbs absent but usual position marked by
an irregularity in a single scale on each side; 22 scales on neck;
22 on anterior part of body; 20 at exact midbody; reduced to 18 a
centimeter anterior to anus; no hind limbs, the position marked by
some irregularities in the scales but no tubercle or remnant of the
limb can be discerned; 121 scale rows on dorsum from nuchal to
above anus; 125 from postmental to anal shield; 95 ventral scales
from anus to tip of tail (complete) .

Color. Ground color brownish fawn, each scale with a semilunar
spot of dark brown on its extreme anterior part and thus covered
by the preceding scale, the spots not or but rarely contiguous ; spots
forming longitudinal rows as well as diagonal rows. An indefinite
blackish brown area above eye, continued down across the preocular.

518 The University Science Bulletin

Measurements. Total length, 130 mm.; snout to vent, 80 mm.;
tail, 50 mm.; width of head (greatest), 4.15 mm.; length of head to
posterior edge of parietals, 5.6 mm.; width of body, 5 mm.

Remarks. At least two species occur in the flat northern lowlands.
These are Nessia sarasinorum and the present species. The charac-
ter of the head scales (especially in regard to the loreals) differs in
numerous regards from Nessia layardi with which it agrees in ab-
sence of limbs and ear-opening.

The species is named for Mr. P. E. P. Deraniyagala, noted Cey-
lonese herpetologist, and Director of Museums, Ceylon.

THE UNIVEESITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXIII, Pt. II] March 20, 1950 [No. 14

The Snakes of Ceylon

BY

Edward H. Taylor,

Department of Zoology, University of Kansas

Abstract: The report is based primarily on the Ceylonese snakes in the
collection of the United States National Museum, and those in the Edward
H. Taylor-Hobart M. Smith collection at Lawrence, Kansas. The paper also
includes a few specimens recently received in exchange from Dr. W. C. Osman-
Hill and now in the University of Kansas Collection.

Two new species, Ahaetulla oliveri, and Lycodon osmanhilli are described.
Scale data are included for most of the specimens in the collections.

The island of Ceylon is situated southeast of the tip of the
Indian peninsula at a distance of about 40 miles from the mainland.
The distance between is partially bridged by a small peninsula and
several islands, commonly called Adam's Bridge. The peninsula
pushes off from the mainland not far from the mouth of the Vaigai
river and is, seemingly, a part of the delta of that river. Separated
from the peninsula by the narrow Pamban Pass is Rameswaram
island, several miles long. From the Ceylon side six islands stretch
out towards India separated from each other by narrow straits
usually less than two miles wide and from Rameswaram Island by
a gap of approximately 20 miles of very shallow water, so shallow
in fact that from the air the bottom can be seen for much, if not
all the distance.

W. T. Blanford* states that in Eocene and Miocene time the
western coast of India lay farther westward than it does at present
but that this western extension has disappeared through sub-
sidence leaving several island groups. R. D. Oldham* has postu-
lated that the subsidence was accompanied by the formation of a

• H. B. Medlicott and W. T. Blanford. A Manual of the Geology of India, 1st ed., 1879-
2d ed., 1893 (by R. D. Oldham).

(519)

520 The University Science Bulletin

great fault along the west coast. Depths of 2,000 fathoms are now
knowTi between the mainland of India and these islands, the Mal-
dive, Laccadive and Chagos archipelagoes, which were presumably
a part of the mainland formerly.

There is evidence f of a more recent (possibly Pleistocene) sub-
sidence. Excavations for a dock in Bombay have revealed a sub-
merged forest, some of the stumps being thirty feet below sea level.

Moreover the reptilian fauna of Ceylon is similar enough to south-
ern India to warrant a postulation that the separation of Ceylon
from the continent has taken place in relatively recent times, and
that the bulk of the fauna reached Ceylon at a time when the con-
nection was present, and at a much later time than Eocene or
Miocene.

That Ceylon has been affected by subsidence in this general area
would seem quite likely. A very small elevation of the land now
would again bring Ceylon in contact with India ; or even a lowering
of the sea surface by the piling up of glacial ice on the land might
affect sea levels to such an extent as to bring about its union with
the mainland. If subsidence has been a slow process corals might
keep the isthmus built up to near sea level much as occurs in the
formation of atolls. I do not know whether Adams bridge shows
such a condition or not.

The serpent fauna of Ceylon bears a close relationship to that of
the neighboring part of India. This is true for practically all
genera known in Ceylon ; but the fauna is far from being identical.
Many species have been reported as occurring in both India and
Ceylon and are or have been regarded as being identical subspecifi-
cally. However, some of the more recent work of Wall (1921) , Smith
(1943) and Deraniyagala (1948) recognizes subspecific differences
in certain forms previously regarded as identical by Boulenger
(1890).

The fact remains that the matter of subspecific and specific dif-
ferences in the faunas of the two areas is still not clearly defined.
Colonel Wall, who has written on Ceylon faunae, but whose primary
field of interest was the Indian serpent fauna, all too often in his
Ophidia Taprobanica utilized Indian specimens for his descriptions,
and as a basis for data given on the species. In consequence it is
often impossible to judge from his work the exact status of a Ceylon
form, and the extent to which it differs from that of the Indian
mainland. I venture to suggest that one of the most important

t R. B. S. Sewell, Mem. Asia Soc. Bengal, 1935.

Taylor: The Snakes of Ceylon 521

contributions one might make to the herpetology of Ceylon would be
accumulation of considerable series of specimens from many parts
of the Island, as well as from the adjoining southern part of India.
Then, and only then, will it be possible to deteraiine the true re-
lationships and the taxonomic status of many of the forms here
treated. I anticipate that a study of such material would reveal
many other forms differing from Indian species; and furthermore
one would expect to find certain Indian species represented by two
or more subspecies in Ceylon.

There is some evidence of relationship indicated between faunas
of Madagascar and Ceylon. This has been usually explained either
by postulating the presence of a continental land mass connecting
Ceylon, South India and Madagascar most of which is now lost by
subsidence; or by postulating "floating continents." The evidence
based on the serpent fauna is indeed small. However the presence
of a species of Sibynophis in Ceylon and two in Madagascar (one
also in the Comoro Islands), and their presumed absence in Africa
needs some special explanation. This is made even more evident
if one considers the close relationship between the Ceylon lizard
genus, Nessia, and the Madagascar Acontias. Similar relationships
between Ceylon and Madagascar are evident in certain other verte-
brate groups.

A careful examination of the serpent fauna indicates differences
that suggest early arrivals in the country and late arrivals, al-
though the actual or approximate geologic time might be impossible
to determine. This is based primarily on degree or extent of evolu-
tion. One might regard Typhlopidae and Uropeltidae, each with
several endemic species, as belonging in a class with the early
arrivals. Each of these must have undergone a long evolution on
the island, despite the fact that certain widespread forms of the
families occur both in India and Ceylon. Of the very large group
comprising the Colubridae I would regard the Sibynophinae, with
the genus Sibynophis, and Aspidura of the Colubrinae, as two of
the older genera in Ceylon.

There are five genera presumably endemic in Ceylon: Pseudo-
typhlops belonging to the Uropeltidae with one species; Cercaspis
with a single species, having highly modified vertebrae ; Haplocerciis
with one species; Balenophis, an opisthoglyph with specialized
nuchal glands; and Aspidura with five species (one of which has
been reported from the Maldive Archipelago to the west of Ceylon).

522 The University Science Bulletin

The last four mentioned forms are members of the subfamily
Colubrinae.

From the evidence at hand the presumably ancient snakes com-
prising the Anillidae and Boidae have had their evolution elsewhere
and only the end products of their evolution persist. These may
very probably have reached Ceylon at relatively late periods after
the maximum of their specific plasticity had passed. At least the
present evidence points to little evolution of these snakes in the
islands. However Deraniyagala considers Python molurus worthy
of subspecific recognition and names the Ceylon form, P. m. pimbura,
chiefly on the character of the body pattern (a figure is included
here showing both dorsal and lateral markings as well as the dorsal
squamation of the head) (PI. XIII, fig. 1).

Among the other families the Viperidae is represented by three
forms, two of which are regarded as identical to the Indian forms
{Vipera russelli and Echis carinatus). These possibly are of recent
arrival since E. carinatus is found only in the extreme northern part
and V. russelli apparently is in lowland chiefly ; this despite the fact
that in India it has been reported up to elevations of 7,000 ft. On
the other hand the pit vipers {T rimer esurus and Agkistrodon) seem
to have undergone considerable evolution in the islands and may
have preceded the two genera mentioned previously by considerable
time. One typical T rimer esurus is endemic at high elevation in
Ceylon, and the genus Agkistrodon has itself undergone change and
one endemic form is recognized.

Representing the Elapidae are three genera: Bungarus with two
species (one endemic) ; Naja with presumably one subspecific rep-
resentative; and Callophis with a form probably subspecifically
distinct from Indian forms (here not so regarded).

The table which follows compares the fauna of southern India
and Ceylon by genera: The endemic genera in southern India are
six, while endemic genera in Ceylon number five (in one of these,
a species is represented also in the Maldive Archipelago but not in
India). There are 74 species in Ceylon, of which 37 are presumably
endemic. In southern India 102 species are known.

Taylor: The Snakes of Ceylon

523

Fig. 1. Map of Ceylon showing provinces.

524

The University Science Bulletin

Table of Snake Genera for Peninsular India and Cetlon Excx-usive of the

Sea Snakes Hydrophiidae

Ceylon

Typhlops 10

Uropeltis 2

Melanophidiiim

Platyplectrurus

Teretrurus

Plectiiirus

Rhinophis 7

Pseudotyphlops 1

Cylindrophis 1

Python 1

Eryx 1

Acrochordus 1

Sibynophis 1

Elaphe 1

Ptyas 1

Coluber 1

Liopeltis 1

Oligodon 4

Ahaetulla 4

Chrysopelea 2

Lycodon 3

Cercaspis 1

Dryocalamus 2

Natrix 3

Balanophis 1

Macropisthodon 1

Atretium 1

Rhabdops

Aspidura 5

Haplocercus 1

Xylophus

Boiga 5

Dryophis 2

Enhydrus

Hurria 1

Gerardia 1

Bunganis 2

Callophis 1

Naja 1

Vipera 1

Echis 1

Agkistrodon 2

Trimeresrus 1

Ceylon
endemic
species

8
2

7
1
1

4
1

2
2

India

6
14
3
2
2
4
3

2
2

2

9
5
2

4

2
4

"i

1

1

(iMaldiveA.)

2
3
4
1
1
1
2
3
2
2
1
2
4

Taylor: The Snakes of Ceylon 525

Family TYPHLOPIDAE

A single genus, Typhlops, is represented in Ceylon, no less than
ten species being present.

Genus Typhlops Oppel

Typhlops Oppel, Die Ordnungen, Familien und Gattungen der Reptilian . . ., 1811,
p. 54.

Genotype, Anguis lumbricalis.

The snakes of the genus Typhlops that are known to occur in

Ceylon have been treated in a special paper.* However, in order

that the listing here may be complete, I have also included these

species.

Key to the Species of Typhlops in Cbylon

1. Scales in 18 longitudinal rows about body ; nasal variable 2

Scales in 20 or 22 rows about body ; nasal divided 4

2. Nasal incompletely divided, the suture to 2d labial; no subocular; small terminal

tail spine; eye distinct; 330-360 transverse scale rows; length 285 mm.;

blackish brown above, paler below T. porrectus

Nasal completely divided, the suture to 2d labial ; a subocular present ; no ter-
minal tail spine ; eye usually not visible 3

3. Upper (posterior) nasals separated behind rostral; 330-360 transverse scale rows

on body, brown above, paler below; length 140 mm.; head partly or almost

entirely cream T. mirus

Upper (posterior) nasals forming a median suture behind the rostral ; 330 trans-
verse scale rows on body; brown above, yellowish white below; length 140
mm T. ceylonicus

4. Scales in 20 longitudinal rows about body 5

Scales in 22 longitudinal rows about body ; eye distinct ; nasal suture to 2d

labial; (260-280 scale rows, said to agree with jerdoni). Black above, whitish
below, the colors meeting in a clear line of demarcation; length 130 mm.,

T. leucomelas

5. Nasal suture goes to the preocular 6

Nasal suture goes to second labial 9

6. Transverse rows ot scales on body more than 280 8

Transverse scale rows on body less than 275 7

7. Transverse scale rows, 229-261 on body; subsquamous glands on head forming

distinct pattern ; eyo distinct ; underside of head largely cream ; part of rostral

visible above shorter ; length 130 mm T. lankaensis

Transverse scale rows on body, 245; subsquamous glands on head not visible; eye
rather indistinct ; part of rostral visible above much elongate ; snout somewhat
truncate at tip ; dull violet to lavender; length 111 mm T. violaceus

8. Transverse scale rows on body, 290-330. Pattern of subquamous glands distinct;

eye normally distinct; blackish brown above, paler below; length 170 mm.,

T. braminus

9. Body more attenuated ; transverse scale rows on body above 290 10

Body less attenuated; transverse scale rows 261-273; body brownish above, the

pigment becoming less on sides ; venter cream save for a pigmented band on
throat; eye distinct; width into body length, about 31 times; length 107 mm.,

T. malcolmi
10. Transverse scale rows, 298-326; width of the body into length about 43 to 56
times ; light brown above, lighter below. Scales bordering mouth cream ;
snout rounded anteriorly ; upper nasal one-third larger than lower ; 3d labial

about half area of fourth ; length 112 mm T. tenebrarum

Transverse scale rows 295 ; body width into total length about 60 times ; head
truncate, subsquamous glands not or scarcely discernible; length 90 mm.,

T. veddae

* Taylor, E. H. Comments on Ceylonese Snakes of the Genus Typhlops with descriptions
of new species. Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947, pp. 283, 298, figs. 1-3.

526 The University Science Bulletin

Typhlops porrectus Stoliczka

Typhlops porrectus Stoliczka, Journ. Asiat. Soc. Bengal, vol. 40, 1871, p. 426, pi. 25, figs.
1-4 (type locality, Bengal, India); Mehely, Termes. Fiizetek, vol. 20, 1897, p. 62 (Kala-wewa,
Ceylon) ; Smith, Fauna British India, Ceylon and Burma including the whole Indo-Chinese
Subregion; Reptilia and Amphibia, vol. 3, 1943, p. 46 (Punduloya, Ceylon); Taylor, Univ.
Kansas Sci. Bull., vol. 31, pt. 2, 1947, p. 284.

This species is widespread in India but apparently it is uncom-
mon in Ceylon. Two records, that of Mehely in 1897, and that of
Smith in 1943 are the only specimens that have been reported.

Typhlops mirus Jan

Typhlops mirus Jan, Iconographie Generals des Ofidiens, livr. 1, 1860, p. 9, pis. 5 and 6,
fig. 7 (type locality, Ceylon); Giinther, Reptiles of British India, 1864, p. 176, pi. 16, fig. H;
Theobald, Descriptive Catalogue of the Reptiles of British India, 1876, p. 126; Boulenger,
The Fauna of British India including Ceylon and Burma; Reptilia and Batrachia, 1890, p. 240;
and Catalogue of the Snakes in the British Museum (Natural History), vol. 1, 1893, p. 52,
Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921, pp. 7-9, fig. 1; Journ. Bombay
Nat. Hist. Soc, 29, 1923, p. 348; Smith, The Fauna of British India including Ceylon and
Burma; Reptilia and Amphibia, vol. 3, Serpentes, Dec, 1943, p. 55 (Ceylon. Known definitely
from Peradeniya); Taylor, Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947, pp. 284-285.

Two specimens, EHT-HMS No. 30094, from Peradeniya, Ceylon,
and U. S. N. M. No. 56287 "Ceylon" are in the collection. The
transverse scale rows of these specimens are 317 and 298, respec-
tively.

Typhlops celyonicus Smith

Typhlops mirus Wall (part), Ophidia Taprobanica or the Snakes of Ceylon, Colombo, 1923,
p. 7. (In one place, in a table, the nasals are reported as meeting behind rostral. At the
bottom of the page he states "sometimes in contact behind the rostral.")

Typhlops ceylonicus Smith, Fauna of British India, Ceylon and Burma, including the whole
of the Indo-Chmese Subregion; Reptilia and Amphibia, vol. 3, Serpentes, Dec, 1943, pp. 55-56
(type locality, Peradeniya, Ceylon); Taylor, Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947,
pp. 284-285.

The type of this species is unique.

Typhlops braminus (Daudin)

Eryx braminus Daudin, Histoire naturelle generate et particuliere des Reptiles, vol. 7, year
11 (:= 1803), pp. 279-280. Based on Russell's Rondou-Taloulou-pam in An account of Indian
Serpents collected on the Coast of Coromandel, vol. 1, p. 48, pi. 43 (type locality, Viza-
gapatam, India).

Typhlops braminus Boulenger, Catalogue of the Snakes in the British Museum (Natural
History), vol. 1, 1893, p. 16; Wall, Ophidia Taprobanica or the Snakes of Ceylon; Colombo,
1921, pp. 7, 9-13 (figures apparently represent another species); Smith, The Fauna of British
India, Ceylon and Burma, including the whole of the Indo-C!hinese Subregion; Reptilia and
Amphibia, vol. 3, Serpentes, Dec, 1943, pp. 46-48, fig. 14, head; Taylor, Univ. Kansas Sci.
Bull., vol. 31, pt. 2, 1947, pp. 290-291.

Two specimens are in the U. S. N. M. collection, Nos. 120336,
120337 from Clodagh Estate, Rattota, Matale District, and a single
specimen is in the EHT-HMS collection from Peradliniya, Central
Province. The ventral scales from mental to vent are, respectively,
297, 325, and 317.

Taylor: The Snakes of Ceylon 527

Typhlops malcolmi Taylor

Typhlops malcolmi Taylor, Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947, pp. 291-292 (type
locality, 12 mi. N Trinconnalee, Ceylon).

The species is known from the type and a single topotypic para-
type specimen, both in the EHT-HMS collection.

A small bicolored snake having 20 longitudinal scale rows; 261-
273 transverse scale rows around body; fourth labial twice as large
as third, notched behind; a single postocular present; brown above
and cream below wdth a brownish pigmented band crossing throat.
Glands on head form a distinct, lighter pattern.

Typhlops violaceus Taylor

Typhlops violaceus Taylor, Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947, pp. 289-290 (type
locality, 12 mi. N Trincomalee, Ceylon).

This species is known from the type specimen (EHT-HMS collec-
tion) onl}', and may be diagnosed as follows: A small Typhlops
with eyes dim, but discernible; the nasal completely divided, the
nostril lateral but not visible above, the suture dividing nasal touch-
ing the preocular far from the labial; 20 longitudinal scale rows;
none or only a very small terminal spine ; 245 transverse scale rows
en body; body width in total length about 31 times. Color, dull
violet to lavender, almost the same above and below.

Typhlops veddae Taylor

Typhlops veddae Taylor, Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947, pp. 294-296, figs. 3,
A-B (type locality, 12 mi. N. Trincomalee, Ceylon).

The unique type specimen is in the EHT-HMS collection. The
diagnostic characters are as follows : A very slender Typhlops hav-
ing 20 scale rows, no subocular, the nasal suture to the second labial,
the rostral short failing to reach back to eye level by a considerable
distance; head somewhat narrowed, and truncate anteriorly; the
transverse scale rows about body 295; 14 caudal rows; body width
in length about 60 times.

The species, judging by its very slender habitus, and the large
number of transverse scale rows is related to Typhlops braminus,
and to Typhlops tenebrarum, described recently. It differs from the
former in color, characteristic pattern of glands, and in having the
nasal suture touch the labial rather than the preocular. The shape
of the head and particularly the shape of the snout, is quite differ-
ent; the eye is dim, and the diameter of the body is contained in its
length 60 instead of "30-45" times, and the tail is proportionally
longer.

528 The University Science Bulletin

The type was taken from decaying wood debris, in second growth

forest.

Typhlops leucomelas Boulenger

Typhlops leucomelas Boulenger, The Fauna of British India including Ceylon and Burma;
Reptilia and Batrachia ; London, 1890, pp. 237-238 (type locality, Haycock Mountain, 40 mi.
from Galle, Ceylon, 2,000 ft. elevation) ; and Catalogue of the Snakes in the British Museum
(Natural History), vol. 1, 1893, pp. 18-19, pi. 1, fig. 4; Wall, Ophidia Taprobanica or the
Snakes of Ceylon, 1921, pp. 13-15, fig. 4; and Spolia Zeylanica, vol. 12, 1922, p. 253; and
Journ. Bombay Nat. Hist. Soc, 29, 1923, p. 350; Smith, The Fauna of British India, Ceylon
and Burma including the whole of the Indo-Chinese Subregion; Reptilia and Amphibia, vol. 3,
Serpentes, Dec, 1943, p. 50; Taylor, Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947, pp. 286-287.

No specimens of this species are in the collection.
Typhlops lankaensis Taylor

Typhlops lankaensis Taylor, Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947, pp. 287-289, figs.
1, A and B (type locality, 12 mi. N Trincomalee, Ceylon).

This form occurs commonly in the Trincomalee area. The type
and 24 paratypes are in the EHT-HMS collection. The diagnostic
characters follow:

Head oval, seen from above; suture, dividing nasal completely,
reaches to preocular; 20 scale rows about body; rostral somewhat
less than one third greatest width of head; transverse scale rows
about body, 229-261; generally brown to gray-brown above with
very dim longitudinal dorsal lines, the median most distinct; under
side of head very largely cream color.

Typhlops tenehrarum Taylor

Typhlops tenehrarum Taylor, Univ. Kansas Sci. Bull., vol. 31, pt. 2, 1947, pp. 292-294,
figs. 2, A-C (type locality, 12 mi. N. rrinconialee, Ceylon).

The type and three paratypes are in the EHT-HMS collection.
The diagnostic characteristics of the species are as follows: A very
slender Typhlops with the nasal suture completely dividing the
nasal and reaching the second labial ; width contained in length from
about 43 to 56 times; number of transverse scale rows about body
from about 298 to 326 rows ; 12 to 14 on tail ; 20 longitudinal scale
rows about body; light brown above, lighter below and on sides;
scales bordering the mouth cream color.

Family UROPELTIDAE

Of this family the genera Uropeltis, Rhinophis, and Pseudotyph-
lops occur in Ceylon. Uropeltis is represented by two endemic
species (a third form bears the name ceylonicus but apparently it
is an Indian snake) . Rhinophis by eight species and Pseudotyphlops
by one. Thus eleven of the 45 species known are Ceylonese. One

Taylor: The Snakes of Ceylon 520

genus, Pseudotypldop)<, and eight of the eleven known Rhinophis are
endemic in Ceylon. However, of the species of Uropeltis, 20 of the
22 known species are Indian in distribution.

Key to the Ceylonese Genera of Uropeltidae

1. Tail usually obliquely truncate, the truncate portion covered with thickened dif-

ferentiated scales; terminal caudal scute ending in a transverse ridge, or two

points side by side Uropeltis

Tail not obliquely truncate 2

2. Tail ending in a convex or flattened, rounded, rugose shield Rhinophig

End of tail with a large, subcircular, fiat, spinose shield above Pseudotyphlops

Genus Rhinophis Hempricht

Rhinophis Hempricht, Grimdrlss der Naturgeschichte, 1820, p. 119.

Genotype, Tylops oryrhynchus ; Wagler, Natiirliches System der Amphibien. 1830, ]). 10.'>.

The genus is represented by eight species in Ceylon. Three others
occur in India.

Key to the Species of Rhinophis in Ceylon

1. Rostral lacking a lidge 2

Rostral more or less distinctly ridged 3

2. Ventrals 148-168; dark brown, each scale below with a yellow spot or niartiin ;

yellow vertical spots on each side on anterior part of body usually connected

by a lateial stripe ; length 370 nun blytlii

Ventrals 173-191 ; brown above, uniform or each scale with a white area or mar-
gin, more extensive below ; light spots may be present, or bars on sides ; a
ring around base of tail ; length SOO mm clrummoiidhayi

3. Rostral ecjual or less than half the length of the headshiekis; rostral obtusely

ridged 4

Rostral more than half the length of the headshiekis; strongly ridged 5

4. Ventrals 180-204; blackish brown, the scales with lighter margins; a series of

cream spots along sides homolepis

Ventrals 153-182; brown above and lielow with the scales somewhat lighter mar-
gined ; no lateral creani spots pliilippinus

0. L'niform brown above and below; ventrals 211-227; rostral reddish.... oiyrhynrlius

Not uniform brown above and below 6

G. A black vertebral line between two light lines 7

No black vertebral line between two light lines 8

7. Ventrals 236-246 ; diameter in length 46 times pimctatus

Ventrals 281 ; diameter in length 76 times porrectus

8. A broad orange stripe dorsally, bearing black blotches; ventrals 238 dorsimaciilatiis

Rhinophis blijthi (Kelaart)

Hliiiiophis blytlii Kelaart, Prodromus Fauna Zeylanica, vol. 2, 1854, p. 14 (type locality.
Mountains of Ceylon); Smith, Fauna of British India . . ., Reptilia and .\mphibia, vol. 3,
1943, pp. 88-89.

No specimens in the collection.

Rhinophis dnimmondhayi Wall

Rhinophis dnnnmondhiiyi Wall, Ophidia Tapiobanica or the Snakes of Ceylon, 1921, iip.
43-44 (type locality, U\a Patnas, Ceylon).

No specimens in the collection.
33—90

530 The University Science Bulletin

Ehinophis porrectus "Wall

Rhinophis porrectus Wall, Ophidia Taprohanica or the Snakes of Ceylon. 1921, pp. 3.t-,'?6
(type lop.ility, "on road between Puttalani and Chilaw").

No specimens in the collection.

Smith has placed this form in the synonymy of Rhinophis punc-
tatus. I cannot agree since the characters are too striking to war-
rant such an association. The ventral count is 281, 35 above the
highest known count for pimctatus, and the body is slenderer, the
diameter in total length about 76 times while Smith's specimen of
R. punctatus has the diameter in total length, 47.5 times.

The length is 355 mm. The head is black brown, while the tip of
snout is dull orange. A narrow blackish brown vertebral line passes
from the nape to near the end of the body occupying the medial
part of the vertebral scale row. This is bordered by whitish stripes.
The caudal shield is dull orange with a subterminal dark mark. The
species is known from Northwest Province.

Rhinophis punctatus Miiller

Rhinophis punclafus jVIuller, Tiedemann and Treviranus Zeitschrift fiir Physiol., vol. 3,
p. 1832, pp. 248-249, pi. XXI. figs. 1-3 (cranimn), pl. 22, figs 1, a-c (head), d-f (tail) (type
locality, Ceylon); Peters. De Serpentiim familia Uropeltaceoruni, 1861, pp. 12-13, pl. 2, fig. 3;
Smith, Fauna of British India . . ., Reptilia and Amphibia, vol. 3, 1943, p. 92 (in part).

No s]jecimen is in the collection. The characters given in the key
will suffice to separate the form from others known in Ceylon. The
species is reported from Central Province: Kandy and Peradeniya.

Rhinophis philippinvs (Cuvier)

Typhlops philippinus Cuvier, Regne .\nimal, 2d ed., vol. 2 1829, p. 74.

Rhinophis philippinus Miiller, Tiedemann's und Treviranus' Zeitschrift fiir Physiol., vol. 4,
1832, p. 249; Dumeril, Bibron and Dumeril, Erpetologie Generale, vol. 7, 1854, p. 154,
Atlas, pl. 59, fig. 1; Peters, De Serpentum familia Uropeltaceorum. Berlin, 1861, pp. 15-16;
Gunther, Reptiles of British India, 1864, p. 184; Smith, Fauna of British India . . .,
Reptilia and Batrachia vol. 3, 1943, p. 91.

Rhinophis planireps Peters, De Serpentum familia Uropeltaceorum, Berlin, 1861, p. 17, pl. 1,
fig. 9.

A specimen in the EHT-HMS collection (No. 30387) from Pera-
deniya and U. S. Nat. Mus. No. 56402, "Ceylon," are referred to
this species. The first is uniform brownish, the scales showing some-
what lighter edges, the ventral scale rows having the light edges a
little wider than the dorsals. The under side of the snout, the lower
labials and most of the two anal plates are cream. The top of the
head is convex, not flattened, and the rostral length above is some-
what more than one third of the shielded part of head, but much
less than one half. There are four supralabials and three infra-
labials. An azygous preanal, triangular in shape is present. The

Taylor: The Snakes of Ceylon 531

anal is large and divided, followed by a divided postanal only a
little smaller than anal. This is followed by five subcaudals all of
which are undivided except the fifth. The shield on the tail is as
long as the shielded part of the head. The ventrals are 155 in this
male specimen; subcaudals as described, five.

No. 56402 is blackish brown with light edges on the scales. The
under side of the labials and rostral, and a part of the divided anal
scale are cream. The ventrals are 174 5 ; subcaudals: one divided
postanal followed by a divided subcaudal and one single scale to-
talling three.

Two specimens, U.K.M.N.H. Nos. 21433 "Ceylon," and 21434
Polgahaivela, Ceylon, have the following characters respectively:
total length, 183 mm., 176 mm.; tail, 5, 7; ventrals, 177, 179; sub-
caudals, 3, 6; scale formulae, 19, 17, 17, 17; 19, 17, 17, 17; diameter
of body, 8.8 mm., 6.8 mm.

The rostral length is slightly less than its distance from the rostral
to back of frontal in both. In the longer the rostral separates the
prefrontals for less than one half the suture, in the shorter the sep-
aration is more than half the length of the suture.

Rhinophis oxyrhynchus (Schneider)

PI. XII, fig. 2

TypJiliipx ojijrhyiichuK Schneider, Historia AmphiMdiuiu. vol. 2, p. 341 (tj-pe locality,
Ceylon).

Rhinophis oxyrhynchus Henipricht, Gnindriss der Natmgeschichte, 1820, p. 119; Dunieril,
Bihron and Dunieril, Erpetologie Generale, vol. 7, 1854, pp. 156-157 (oxyrhynchus).

Dapatiiaya lankadivana Kelaait. Prodromus Faunae Zeylanicae, vol. 2, 1854, pp. 10-17
(type locality, "Common at Trinconialee and in the Kandyan Province").

Mytilid iiiiiinaculata Gray, Proc. Zool. Soc. London, 1858, p. 2(54. fig. (type locality,
Ceylon).

I collected a specimen, EHT-HMS No. 31256, about 6 miles north
of Trincomalee that is typical in its general characters. The ven-
trals are 216 ; the subcaudals 8 ; the scale rows on neck, 19 ; on the
body 17. A pair of elongate postanal scales are followed by seven
subcaudals. The median dorsals are much widened on tail. The
rostral is distinctly longer than its distance to the back of the
parietals. The length is 329 mm., the tail, 8 mm. The diameter
of the body is 8 mm.

The species is known from Northern and Eastern provinces (in the
north). I collected a specimen crawling in the road at midday 14
miles north of Trincomalee. It escaped from a faulty collecting bag
into a raft while crossing a lagoon and could not be recovered.

532

The University Science Bulletin

Plate XII. Fis. 1. RhinopJnti hojnolepis'H.emYtvich. U. S. N. M. No. 56430;
"Ceylon"; total longth, 243 mm. Fig. 2. Rhinophis oxyrhynchus (Schneider).
EHT-HMS No. 31256; 6 mi. north Tiincomalee, Eastern Province. Ceylon.

Taylor: The Snakes of Ceylon 533

Rhinophis homolepis Hempricht

PI. XII, fig. 1

Rhinophis homolepis Hempricht, Grundriss der Naturgeschichte, 1820, p. 119 (type locality,
Ceylon fide Peters); Peters, De Serpeiitum Familia Uropeltaceorum, 1861, pp. 14-15, pi. 2. fig.
2; Smith, Fauna of British India . . ., Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp.
90-91.

Dapatnaya trcveJyaim Kelaart, Prodromus fauna Zeylanicae, vol. 2, pt. 1, 1S.")4, ji. 17 (type
locality, Kandyan Hills).

Mitylia or Mytilia Gerrardi Gray, Proc. London Zool. Soc, vol. 26, 1858, pp. 58, 2(13, i)l.
13 (type locality, Ceylon).

A specimen in the collection (U. S. Nat. Mus. No. 56430) from
"Ceylon" has the following scale characters: Ventrals 197, some-
what enlarged; 3 siibcaudals ; anal divided; scale formula, 19-17.

The head is diminutive, the shielded part being shorter than the
terminal caudal plate. There are 23 lateral white spots, the largest
of which are on the neck. The postanal shield is white. The species
is known from Central Province, Uva Province, and Sabaragamuwa
Province.

Rhinophis dorsimacuUitus Deraniyagala

Bhinophis dorsi7)iacidattis Derani.s'agala, .loiirn. Bombay Nat. Hist. Soc, Dec, 1!)41, jip.
800-802, pi. and text fig. 1 (t>pe locality, Marichchukate, Northwest Province, Ceylon); Smith,
Fauna of British India . . ., Reptilia and .\mphibia, vol. 3, Serpentes, 1943, p. 526.

This recently described species is known from the two type speci-
mens, which were found in the arid part of Ceylon in the north. It
is characterized by a broad dorsal orange stripe with small dorsal
spots. The total length of the largest specimen is 350 mm. The
ventrals are 238 and the scales are in 17 rows. .

Genus Uropeltis Cuvier

Vropeltis Cuvier, Regne .\nimal, 2d ed., 1829, vol. 2, p. 76 (part).
Cienotype, ceytanicus.

Two forms of the genus occur in Ceylon.

Key to the Species of Uropeltis ix Ceylox

1. \'entrals 141-166; belly black; dark brown above with an irregular lateral stripe,

melanogaster
Ventrals 197-210; bluish gray, with 7 median dorsal rows bearing yellow longi-
tudinal lines made of yellow dots; a lateral series of yellow bars phillipsi

Uropeltis melanogaster (Gray)

Mytilia (Crealiaj ynelanogafter Gray, Proc. Zool. Soc. London, 1858, p. 204, fig. 5 (tj-pe
locality, Ceylon).

Uropeltis melanogaster Smith, Fauna British India . . ., Reptilia and .\mphibia, vol. 3
1943, pp. 86-87.

Four specimens of this species are at hand: U. S. Nat. Mus. Nos.
7134, 56397, 56398, 56400. All are from ''Ceylon." The species is
known from Central Province.

534 The University Science Bulletin

Scale Counts of Uropcltis iticlnnognatcr

Tot

al length

Tail

Number

Sex

Scale rows

Vent nils

Subcaudals

mm.

mm.

7134

2

19-17

167

6

220

7

56397

$

19-17

160

10—4 wide

144

6.8

56398

9

19-17

168

t

200

6.6

56400

$

19-17

160

9—6 wide

145

7

[Ventrals counted from mental. J

The specimens are soniewliat variable in color. No. 56398 is
blackish brown, the venter even darker but bordered by an extremely
ragged cream line, sometimes discontinuous, varying in width and
distinctness. No. 7134 is brown, the light stripes less ragged.

The two males are light brown and the ventral surfaces are largely
yellowish cream with occasional brownish scales. The ventrals
range from 141 to 168 (counting from the mental).

Uropeltis phillipsi (Nicholls)

Silybura phillipsi Nicholls, Ceylon Jouin. Sci., B, vol. 12, 1929, p. 153; and idem, D, II,
1929, p. 97 (type locality, Menakanda Group, E Matale Hills, Ceylon).

Uropeltis phillipsi Smith, Fauna British India . . ., Reptilia and Amphibia, vol. 3,
1943, p. 81.

This form is known only from the type locality and Mouskandy
Hills, Gammadura, according to Smith.

Genus Pseudotyphlops Schlegel

Pseudo-fyplilops (in part) Schlegel, Abbildungen neuer oder unvollstiindig bekannter Amplii-
bien, 1838, p. 40 (type philippiims \fide M. Smith, loc. cit.]).

A single species known. It is endemic in Ceylon.
Pseudotyphlops philippinus (Cuvier)

Uropeltis philippinus Cuvier, Regne Animal, 2cl ed., vol. 2, 1829. )). 74 (type locality,
"Phillippines" ex errore); Miiller, Tiedemanii and Treviranus, Zeitsch. fiir Physiol., vol. 4, pi.
22, figs. 2-3 Gervais Guerin Mag. Zool., 1837, CI. 3, pi. 13; Dumeril, Bibron and Dumeril,
Erpetologie Generale, vol. 7, 1854, pp. 161-103, Atlas, pi. 59, fig. 2 (see considerable synonj-my) ;
Peters, De Serpentum Familia Uropeltaceorum, Berolini, 1801, p. 20.

Pseudotyphlops pliilippinus Schlegel. Al)bildungen neuer oder unvollstiindig bekannter Am-
phibien, 1838, p. 44; Smitli, Fauna British India . . ., Reptilia and Amphibia, vol. 3, 1943.
pp. 93-94, fig. 27.

Uropeltis grandis Kelaart, Prodromus Famiae Zeylanicae, vol. 2, 1854, p. 15 (type locality.
Kerinday, near Matura, South Province, Ceylon).

Uropeltis saffragamtis Kelaart, Prodromus Faimae Zeylanicae, vol. 2, 1854, )). 15 (t\pe lo-
cality, Ratnapoora near Adams Peak. Ce>lon).

Uropeltis pardialis Kelaart, Prodromus Faunae Zeylanicae, vol. 2, 1854, p. l(i (ty|)e localit.v,
Matura, Ceylon).

The collection contains no specimens belonging to this species. It
is a large species of the family, reaching a length of 285 mm. and a
diameter of 22 mm., the latter measurement being much larger than
that of any other member of the family. The species is dark brown
to blackish above, the young having yellow spots. Yellow beneath,
the young with dark brown spots on venter.

A lowland form for the most part.

Taylor: The Snakes of Ceylon 535

Family ANILIIDAE
Genus Cylindrophis Wagler

Cylindrophis Wagler, Icon. Aniphib., 1828, p. 5.
Genotype, resplendens.

One species occurs in Ceylon.

Cylindrophis maculatus (Linnaeus)

Atiguis 7naculata Linnaeus, Museum Regis Adolphi Friderici. p. 21, pi. 21, fig. 3 (type
locality, "America"); and Systema Naturae, vol. 1, 10th ed., 1758, p. 228.

Cylindrophis maculatus Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921, pp. 18-
21, figs. 5-6.

The species is said to be common in Ceylon in the plains and in
the hills at low elevations. There are no specimens in the collection.

Family BOIDAE
Subfamily Pythoninae
Two genera, each represented by a single species, occur in Ceylon.
Key to the Gener.a of the Boid.ae in Ceylon

Teeth on the premaxilla ; anterior labials and rostral pitted Python

No premaxillary teeth ; rostral and labials not pitted Eryx

Genus Python Daudin

Python Daudin, Mag. Encycl., Mar. 1803 (an 8), p. 434.
Genotype, Python molurus.

Python molurus Pimbura (Deraniyagala)

PI. Xlll. fig. 1
Python molurus pimbura Deraniyagala, Spolia Zeylanica, vol. 24, pt. 2, 1945. p. 105
(Colombo, Ceylon).

EHT-HMS Nos. 31258 yg.; 31259 yg.; 31260 (head only).

The presence of Python molurus has long been known in Ceylon.
It is one of the more common snakes of the country. Two specimens
were found fifteen miles north of Trincomalee on a tiny islet about
one and one half miles offshore. One was dead beside a small rock
pool of fresh water having been recently killed, presumably by
fishermen. The second specimen was found submerged in another
fresh water pool containing much algae. The tip of the snout ex-
tended above the surface amid the algae, near the edge. Since the
dead specimen had recently fed on a wood pigeon, it was suspected
that the snakes caught their prey as the birds drank in the fresh
water pools. This was the only species of land snake obtained on
the island. Numerous land birds were accustomed to fly over to
the island daily and the snake population was assured of a regular
food supply about the fresh water pools in the rocks.

Scale counts of Nos. 31258 and 31259 respectively: Ventrals 248,
233; anal 1-1; caudals 61, 63; scale formul-a 55-64-41, 55-59-38.

Platk XIII. Fig. 1. Pi/lltoti niolurn-^ pl))>burn Deraniyagala. EHT-HMS
No. 31258, yg.; 12 miles north Tiincomalee, Eastorn Province, Ceylon; total
length, 612 mm. Fig. 2. Macropisthodon plumbicolor (Cantor). EHT-HMS
No. 30681 9 ; 50 mi. S Amiradapura, Ceylon; total length, 627 mm. (The
characteristic head and neck markings were visible before the loss of the outer
epidermis).

Taylor: The Snakes of Ceylon 537

Genus Eryx Daiidin

Eryx Daudin, Mag. Eiicycl.. vol. 5, 1803, p. 437.
Genotype, turcicus.

Eryx conicus (Schneider)

Boa cnnica Schneider. Historia Amphibiorum naturalis et literaiia .lena 1799-lSOl, vol. 2,
p. 268; Denkschr. Akad. Miinchen, vol. 7, 1821, p. 119. pi. 4, fig. 2 (based on Rus-sell's Indian
Serpents, vol. 1, p. 4, pi. 4 [Madras]).

Eryx conictis Deraniyagala, Ceylon Journ. Sci., B, vol. 19, 193(), p. 33.5. fig.; Smith, The
Fauna of British Indian. Ceylon and Burma including the whole of the Indo-Chinese sub-
regioii ; Reptilia and Amphib:a. vol. 3, Serpentes, 1943, p. 112-113, fig. 35.

No specimens of the sand boa are in the collections. It is re-
garded as a rare snake in Ceylon.

Family COLUBRIDAE

Four subfamilies of the family Colubridae are represented in Cey-
lon. These are:

Acrochordinae, genus Acrochordus.

Homalopsinae, genus Hurria.

Sibynophiinae, genus Sibynophis.

Colubrinae, genera Elaphe, Ptyas, Coluber, Liopeltis, Oligodon,
Ahaetulla, Crysopelea, Lycodon, Cercaspis, Dryocalamus, Aspidura,
Haplocercus, Dryophis, Boiga, Natrix, Balanophis, Macropisthodon,
and Atretium.

Key to the Subfamilies of Ceylonbse Colubrid.\e

1. Dentary free behind, attached to articulare anteriorly; maxillary teeth 30 to 50.

Head with large symmetrical shields: no grooved teeth Sibynophiinae

Dentary not free behind ; maxillary teeth usually les.s than 35 2

2. Head covered with small granular scales; no widened ventral scales; body covered

with loose skin, eyes more dorsal than lateral ; posterior maxillary teeth not

grooved Acrochordinae

Head covered with enlarged regular shields; teeth grooved or not 3

3. Nostril crescentic, on dorsal surface of the snout; rostral without the normal deep

excavations for tongue protrusion; eye small, directed upward; last two or

three maxillary teeth grooved; salt or fresh water snakes Homalopsinae

Nostril not crescentic, not on dorsal surface of snout ; rostral with excavations for
tongue extrusion; eyes lateral, directed outward; maxillary teeth with or with-
out grooves Colubrinae

Subfamily Acrochordinae

Acroclwrdinac Boulenger, Catalogue of the Snakes in the British Museum, vol. 1, 1893, p.
172 (part); Smith, The Fauna of British India . . ., Reptilia and Amphibia, vol. 3, Ser-
pentes, 1943. p. 131.

M. Smith, loc. cit., has united the two genera Acrochordus and
Chersydrus into the single genus Acrochordus, stating that he did
not regard the presence of the ventral abdominal fold a character
of sufficient importance to warrant the retention of Chersydriis.

538 The University Science Bulletin

Genus Acrochordus Hornstedt

Acrochordus Hornstedt, Abh. Acad. Stockhcilin. ITS", \o\. S, p. 30".
Genotype, Acroohordus javanicus.

Acrochordus granulatus (Schneider)

Hydras granulatus Schneider, Historia Aniphibioruni naturalis et literaria, vol. 1, 1799, p.
243 (type locality, India).

Acroclurrdus gramdatus Shaw, General Zoology, vol. 3, 1802, p. 576, pi. 130; Smith, The
Fauna of British India . . ., Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp. 134-13.5.

No specimens in the collection.

Subfamily Sibynophinae
Two genera are known.

Genus Sibynophis Fitzinger

Sibytwphis Fitzinger, Systema Reptilium, 1843, p. 26.
Genotype, Herpetodryas geminatus Schlegel.

Taylor and Smith* have separated the Mexican and Central Amer-
ican species of this subfamily into a separate genus, Scaphiodonto-
phis.

The range of the genus Sibynophis includes Madagascar, Comoro
Islands, Ceylon and south Asia from India to China. There are
ten valid species known.

Sibynophis siibpunctatus (Dumeril, Bibron and Dumeril)

Pi. XIX, fig. 1

OUgodon suhpunctatutv Dumeril, Bibron and Dumeril, Erpetologie Generale, vol. 7, 18,54,
p. 58 (type locality, "Malabar").

Polyodontophis suhpunctatum Boulenger, Fauna British India including Ceylon and Burma,
Reptilia and Batrachia, 1890, p. 303 (Ceylon); Catalogue of the Snakes in the British Museum,
vol. 1, 1893, pp. 180-187; Abercromby, Spolia Zeylanica, vol. 8, pt. 32, Jan., 1913, pp. 304-
305 (Coloml)o, Matale, Kalutara) ; Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921,
pp. 84-89, fig. 20 (Colombo; Neboda near Kalutara; Puttalam ; Galatura Estate Colombo;
Ratnapura District, 1,000 ft.).

Enicognathus liumberti Jan, Arch. Zool. Anat. Fis., vol. 2, fasc. 2, Mar. 31, 1863, p. 65;
Elenco systematico degli Ofidi, 1863, p. 50 ("Trincomalie, Ceylan") ; LTconographie Generale
des Ophidiens, livr. 16, 1866, pi. 4, fig. 1 ("Trincomalie, Ceylan").

EHT-HMS Nos. 31251-31253, 12 miles north of Trincomalee;
U. S. Nat. Mus. No. 56231.

This small species has long been known from Ceylon. It is prob-
ably not a rare snake but its dimunitive size prevents it appearing
commonly in collections. Two specimens were taken moving about
at night, and one was found concealed below a small log in the day
time.

* Univ. Kansas Sci. Bull., vol. 29, pt. 2, no. 6, pp. 302-304.

Taylor: The Snakes of Ceylon 539

The squamation characters follow: EHT-HMS 31252. Rostral
visible above for a distance equal to half the length of the inter-
nasals; latter, two thirds the length of prefrontals; frontal a third
longer than its distance from snout tip, the sides not angular; pa-
rietals large, their length equal to their distance from rostral ; nasal
divided ; loreal longer than high ; one preocular, two large post-
oculars; two anterior temporals, the lower widely separated from
the postoculars, wedged between the seventh and eighth labial; tem-

1

poral formula 2 -| ; supralabials 9-9, in the following order of

2
size: 1, 5, 4, 3, 2, 6, 7, 8, 9; infralabials 10-10, four touching the

first chinshields ; 2 pairs of chinshields, subequal.

Scale formula, 17-17-17; ventrals 170, anal divided; subcaudals
(divided) 32 (tail with a part missing).

The ventral and subcaudal counts for EHT-HMS Nos. 31234 $
and 31235 j respectively are: 157, 163; 58, 66.

The ventral and subcaudal variation is large in this species.
Smith, loc. cit., gives for the Ceylon and South Indian (south of lat-
itude 14°) specimens, 157-200 ventrals; India north of latitude 18°,
172-215 ventrals. The all-over count of males, 60-70 subcaudals,
for females, 54-63.

Head dark brownish, a cream white spot covering most of each
supralabial; a lighter band between the posterior part of eyes, which
connects with two lighter irregular marks running above eyes and
on canthus to rostral ; a well-defined cream bar on each side of head
crossing ninth supralabial and passing up covering extreme tips of
parietals. These bars separated medianly by a narrow black longitu-
dinal line; behind these bars, four to six scale rows, a transverse
cream band, only partially interrupted mesially. The dorsal color
is browaiish produced by a peppering of pigment on a lighter back-
ground; a median row of darker punctations often outlined in cream;
on the posterior part of the body a line formed of short cream dashes
is discernible on the fourth and (more posteriorly) fifth scale rows.
A dark spot on ends of the ventrals and subcaudals distinct through-
out entire length on males, somewhat less distinct in the female.

The species is confined largely to the lowdand and hill areas in
Ceylon and is seemingly a rare snake.

540 The University Science Bulletin

Key to the Ceylonbse Genera of the Colubrinae

1. All maxillary teeth solid, lacking grooves o

Posterior maxillary teeth (usually two to four) with grooves 2

2. Pupil round 3

Pupil vertical or horizontal 4

3. Solid maxillary teeth 24-26, subequal ; posterior fangs large; head distinct from

neck ; scales strongly keeled save outer row ; ventrals rounded ; a nuchal gland ;

scale rows 19 around middle of body Balanophis

Maxillary teeth 20-22, the last three or four feebly enlarged and grooved ; scales
smooth or feebly keeled ; scales in 17 rows Chrysopelea

4. Pujiil vertical; body more or less compressed; scales smooth, in 19-29 rows, the

vertebral scales often enlarged; 10 to 14 maxillary teeth, followed by two or

, three enlarged grooved fangs Boiya

Pupil horizontal; maxillary teeth 12. the anterior six or seven teeth gradually en-
larged (or last two abruptly enlarged); these followed after an interspace, by a
series of seven small teeth, and these in liim fcijluwcd by two much enlarged
grooved fangs Dryophis

5. Hypapophyses absent on the posterior dorsal vertebrae, the lower surface of

which is smooth or with a very low keel (3

Hypapophyses developed throughout the vertebral column represented by a dis-
tinct crest or keel or a tubercle 15

('). Posterior maxillary teeth longer than anterior 7

Posterior maxillary teeth not longer than anterior 11

7. Pupil round S

Pupil vertical; six to ten maxillary teeth; scales in ]8 or 15 rows, smooth; the

ventrals angulate Dryocalamus

S. Scales in an even number of rows throughout liody, 14-18, smooth or slightly

keeled ; teeth 20-23 ; a presubocular present Zaocys

Scales in odd number of rows or, at least, odd numbers on anterior part of body. . 9
9. Last two maxillary teeth separated by an interval from rest of series; maxillary
teeth 12-18, increasing in size; a subocular (rarely absent); 21 to 23 scale
rows about middle of body ; large snakes of more than a meter length . . . Coluber
Last two maxillary teeth not separated by an interval 10

10. Maxillary teeth 20-28, forming a continuous series; scales often in even numbers

on latter half of body; two or three (rarely four) loieals; large snakes 2

meters or more in length Ptyas

Maxillary teeth 6 to 16, the posterior strongly enlarged and compressed; scales in
13-21 rows; rostral large; scales smooth; small snakes, less than a meter in
length OUgodnn

11. Pupil round; none of the anterior maxillary teeth enlarged, the series not broken

by an interval ; maxillary bone not strongly arched 12

Pupil vertically elliptic; some of the anterior maxillary teeth enlarged and fang-
like; maxillary strongly arched 14

12. Scales in 19-27 rows, with apical pits Elaphe

Scales le.ss than 19 rows ; apical pits present or absent 13

13. Scales 13-17 rows, the vertebrals not enlarged, without apical pits Liopeltis

Scales 13-15, rows disposed in oblique transverse rows, with enlarged vertebrals,

apical pits present Ahaetulla

14. Scales in 15 or 17 rows, smooth or feelily keeled; subcaudals divided Lycodon

Scales in 19 rows, strongly keeled, subcaudals single Cercaspis

15. Posterior maxillary teeth longest; head distinct from neck; loreal jiresent ; one or

two interna.sals 16

Maxillary teeth subeciual ; head not distinct from neck; scales in 15-17 rows: no

loreal ; internasal single 18

10. Two internasp.ls; nostrils lateral; scales in 20-25 rows, strongly keeled 17

One internasal; nostrils directed upward and outward, scaU-s in 19 rows at mid-
body Atretium

17. Maxillary teeth 20 to 35; posterior teeth not fanglike; scales in 15-19 rows. . . Natrix
Maxillary teeth 11 to 18, followed by two very large fangs; scales in 25 to 27

rows Macropisthodon

18. Maxillary teeth, 20-24; scales 15-17 rows; liea\y spines on sides in anal region

in ^ Aspidura

Maxillary teeth 10-12; scales in 17 rows; no lateral spines in anal region. .Haplocerciis

Taylor: The Snakes of Ceylon 541

Genus Elaphe Fitzinger

Elaphe Fitzinger, in Wagler's Icon. Desc. Amiihib., pt. 3, 1828, text to plate 27 (fide
M. Smith).

Genotype, parreysi =: quatunrlineatus.

A single species of the genus occurs in Ceylon.

Elaphe helena (Daudin)

PI. XIV

Coluber helena Daudin, Histoiie Naturelle des Reptiles, vol. 6, year 11 = 1803 (based on
Russell's plate, Hist. Nat. of Indian Coromandel Serpents, pi. 32); Pearless, Spolia Zeylanica,
vol. 6, pt. 21, 1909, pp. 55 (BaduUa, 2,222 ft. elev.); Wall, Ophidia Taprobanica . . .,
1921, pp. 197-203, fig. 42 ("a common up country snake," Colombo).

Cynophis bi.sfri(/atus Gray, Ann. Mag. Nat. Hist., ser. 2, vol. 4, p. 24(> (type locality,
Ceylon).

Elaphe helena Shaw, Shebbeare and Barker, Journ. Darjeeling Nat. Hist. Soc, vol. 14,
1939, p. 78; Smith, The Fauna of British India . . ., Reptilia and Amphibia, vol.' 3,
Serpentes, 1943, pp. 149-152.

Wall, loc cit., suspects that the Colombo record might be ac-
counted for by an importation from the hills. However, I collected
a specimen 12 miles north of Trincomalee at near sea level and an-
other was seen. It was said to be "not rare" in the area.

The specimen EHT-HMS 30680 5 has the following scale char-
acters: Color of head lavender to fawn-brown with a black line
following the parietal suture and a diagonal line behind eye; a
black spot on upper edge of last supralabial. On body, lavender-
brown with a pair of black lines beginning just back of the parietals
and passing back on the neck for about an inch. Here they become
indistinct but can be dimly traced some distance further. A diago-
nal line on the side of the neck beginning five or six scale lengths
back of the jaw angle and terminating on the end of the tenth ven-
tral; a series of indefinite dark blotches follow the lines, the "spots"
consisting largely of dark outlines on the scales contiguous laterally
with a light spot on the fifth and sixth scale rows and one on the
first, second and third scale rows, the two light spots at least par-
tially outlined with black; about one third of the length back of
head the blotches become almost or completely obsolete; but a
broad dark line, scarcely noticeable on the anterior half of the body
becomes more distinct on the latter half of body and tail, covering
most of the second to fifth rows of scales, reduced to a single scale
row on tail. Below uniform yellow cream.

The scale formula is 29-27-21; ventrals 235; anal single; sub-
caudals 86. The known ventral range is from 217-265; subcaudals
73-100, according to Smith, loc. cit.

542

The University Science Bulletin

I'LAiE XIV. FAaphe helena (Daudin) $ . EHT-HMS No. 30680; 12 miles
north of Tiincomalee, Eastern Province, Ceylon; total length, 1,135 mm.

Taylor: The Snakes of Ceylon

543

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544 The University Science Bulletin-

Genus Ptyas Fitzinger

Ptyas Fitzinger, Svr^ter.m Reptilia, 1843, p. 26.

Genotype, Coluber btunieiibachi.

One species occurs in Ceylon. It is a large snake reaching approx-
imately two meters in length when full grown.

Ptyas mucosiis (Linnaeus)

PI. XV. fig. 1

Coluber mucosus Linnaeus, Museum .Adolplii Friderici regis, vol. 1, p. 37, pi. 23; Systema
Naturae, ed. 10, 1758, p. 226 (type locality, India).

Ptija.t mucoxus Giinther. Reptiles of British India, 1864, p. 240.

The collection contains the following specimens of this species:
EHT-HMS Nos. 30696-30701, 12-13 miles north of Trincomalee,
Ceylon; U.S.N.M. No. 56198, Ceylon.

These apparently agree with specimens from the mainland of
India in most characters of squamation and in color markings.

The markings of the anterior part of the body are poorly defined.
The skin between the scales is whitish, which color sometimes en-
croaches on the scales a little. Thus a whitish reticulation is evi-
dent when the scales are distended but only a trace of it is visible
when scales are normally imbricated. The posterior part of body
has numerous transverse black bands, the edges of which are ir-
regular.

The character of the scale rows is unusual in this genus. The
scale rows across the back part of head are 25-27, while the number of
rows on the posterior part is reduced to 14. On the anterior part of
the body, the scale rows are in odd numbers around the body while
in the posterior part of the body they are in even numbers. This
is accomplished by replacing the median single row with paired
scales. In a related genus, Zaocys, this occurs throughout the length
of the body. Even numbers of scale rows is a rare condition in snake
genera. (See preceding page for table of scale data).

Pl.ate XV. Fig. 1. Ftyas mucu.vi.^ (Linnaou8). EHT-HMS No.
30689$ ; 12 miles north Trincomalee, Eastern Province, Ceylon; total
length, 795 iniii. Fig. 2. Natrix piscator (Schneider). EHT-HMS
No. 30730 9 ; 12 miles north of Trincomalee, Eastern Province, Cey-
lon; total length, 745 mm.

Taylor: The ."^nakks of Ceylox

545

C^ '^S-f^'af --,-»■■ _

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34—91

546 The University Science Bulletin

Genus Coluber Linnaeus

Coluber Linnaeus, Systeina Natura, ed. 10, 1758, p. 216.
Genotype, Coluber constrictor.

Only a single species has been recorded for Ceylon.

Coluber fasciolatus Shaw

Coluber faticiplatus Shaw, General Zoology, vol. 3, p. .")2S (Ba.sed on Russell, An Account of
Indian Serpents, vol. 1, 1796, p. 26, pi. 21. India).

Zamenis fasciolatus Wall, Ophidia Taprobanica . . ., 1921, pp. 101-19.T. fig. 41 (.laffna).

No specimen in the collection.

A specimen of this species has been reported by Haly from Jaffna.
Since this record may be due to faulty data, this form must be re-
garded as a doubtful resident of Ceylon.

Genus Liopeltis Fitzinger

Liopeltis Fitzinger, Systema Rt'ptilium, p. 26.
Cienotype. Herpetodryas tricolor Schlegel.

One species occurs in Ceylon.

Liopeltis calamaria (Giinther)

Cyclophis calamaria Giinther, Catalogue of the Colubrine Snakes in the Collection of the
British Museum, 1858, p. 2.50 (type locality, Ceylon).

Liopeltis calamaria Wall, Ophidia Taprobanica . . ., 1921, p. 251. fig.; Smith, Tlie
Fauna of British India . . ., Reptilia and .Amphibia, vol. 3. Serpentes, 1943. pp. 184-185.

No specimen of this species is represented in the collections.

This form may be identified by the following characters: Scale
formula 15-15-15; an elongate undivided nasal scale; supralabials
7-7, infralabials 7-7. The ventrals of Ceylonese specimens vary
between 127-134; the subcaudals between 67-76.

Olive-green dorsally wdth a fine blackish line along the edge of
the fifth and sixth scale rows, the line tending to break up into spots
anteriorly and continuing well on the tail. Another dark line, less
distinct, follows the edges of the third and fourth scale rows. Belly
pale yellow. Some specimens show lighter stripes. Maximum
length, 407 mm.

Genus Olk;odon Boie

Oliqodon Boie, Isis, 1827. p. 519.
Genotype, OUqodon bitorques.

The following five forms of Oligodon are recognized as occurring
in Ceylon: 0. sublineatus, 0. calamarius, 0. taeniolatus fasciatus,
0. t. ceylonicus, and 0. arnensis.

Key to the Form.s of Oligodon in Ceylon

1. Ventral scale.-; with three brown spots forming continuous lines; small paired
dorsal spots sometimes alternating; no palatine teeth, 4-G pterygoid teeth;

ventrals, ^130-144, 2149-100; subcaudals, ^32-30, 9 23-30 sublineatus

Venter lacking distinct lines 2

Taylor: The Snakes of Ceylon 547

2. Venter with squarish white and black spots; a median dorsal white line and 18

narrow cross bars; ventrals, |J 127-138, 5143-152; subcaudals, ^28-34,

5 20-24; palatine and pterygoid almost or completely edentulous calamarius

Venter lacking squarish black and white spots ; few pterygoid teeth 3

Venter white; pterygoid teeth numerous (15-17) arneiisis

3. A moie or less buff species with two broad dark-brown dorsal lines, separated by

a whitish median line; a narrow dark line from mouth to vent; about 43 dim
or diffuse cross bands, each preceded and followed by narrow flecks of white;
venter white or flesh color; ventrals, ^171-175, 5 184-185; subcaudals,

^ 45-51, O 44 taeniolatus ceylonicus

Pale buff or yellowish brown with a median series of rather large, well-defined,
rounded dark spois along back (some may be broken medially); ventrals,

(J 158-159, 5 174 ; subcaudals, ^ 40-40. 5 40 taeniolatus jasciatm

Oligodon arnensis (Shawl

PL XVI, fig. 2; Text fig. 2

Coluber arnensis Shaw, General Zoology, vol. 3, pt. 2, 1802 (type locality, "Country of
Arnee in the East Indies") (Based on Russell's Indian Serpents, vol. 2, 1790, pi. 38, Arni, N.
Areot.).

Oligodon albiventer Giinther, Reptile.s of British India. 1804. p. 213 (near Kandy, Ceylon).

Oligodon arnensis Smith, Fauna ^if British India . . .. Reptilia and Amphibia, vol. 3,
Serpentes. 1943, pp. 225-226, fig. 77.

Oligodon arnensis albiventer Deraniyagala, Ceylon Journ. Sci., ser. B, vol. 20, 1930. p. 89.

A single specimen referable to this species is present in the col-
lection. This is U. S. Nat. Mus. No. 120332, collected at Peradeniya,
Kandy District, Ceylon, by H. G. Deignan.

The ground color is a deep gray brown, with 17 transverse white-
edged black bands, that terminate laterally on the outer scale row.
The venter and most of the outer first row and part of the second
row, pearly white. However there is some slight scattering of pig-
ment on the tM'o scale rows. Between the transverse bands are faint
traces of intercalated spots. These consist of some black flecks on
the sides, and occasionally a few on the back.

A pair of lines run forward from near the second ventral, crossing
the angle of the mouth, and meeting on the frontal (or narrowly fail-
ing to meet ) . Two other lines arise on either side above the ninth ven-
tral and run forward. These meet seven scales back of the parietal.
A narrow band crosses the base of the snout, involves the eyes and
reaches almost to the mouth. Dark flecks are present on the second
and sixth labials.

There are eleven maxillary teeth, small anteriorly, the ninth sud-
denly becoming much larger, followed by two large knifelike teeth.
Eight palatine teeth present followed after an interspace by 17
pterygoid teeth. There are 19 teeth on the mandible.

The ventrals number 180, subcaudals 46. The anal is divided.
The scale formula is 17-17-15. Total length, 317 mm., tail, 47 mm.
The type is said to have 169 ventrals and 50 subcaudals.

548

Till-: L'xivEKsnY Science Bulletin

A B

Fig. 2. Olir/oiloi, amniHls (Shaw), U.S.N.M. No. 120332. A. Dorsal view of
head. B. Lateral view of head (X 10).

There is a strong probability that (Jiinther's name {albiventer)
should be applied to this form. Series of specimens alone can verify
the relationship.

Pl.\te XVI. Fit:. 1. OVujodon tacniolatm ceylonicm^XaW. EHT-HMS
No. 30167 2 ; 12 miles north of Trincomalee, Eastern Province, Ceylon;
total lengtii.'370 nun. Fiji. 2. OUgodon anietmis (Shaw). U.S.N.M. No.
120332 9; Peradeniya, Kandy District, Central Province, Ceylon; total
length, 314 mm.

Taylor: The Snakes of Ceylon

549

-I. V >

550 The University Science Bulletin

Oligodon taeniolatus (Jerdon)

Coronella taeniolatus Jerdon, Journ. Asiat. Soc. Bengal, vol. 22, 1853, p. 528 (type locality,
doubtful, perhaps based, at least in part, on Russell, 1790, pi. 19, p. 24 [V'izagapatam] ).

OUyodon taeniolatux Wall, Opiiidia Taprobanica or the Snakes of Ceylon, 1921, pp. 239-245,
fig. 50.

At least two subspecies occur in Ceylon: 0. taeniolatus fasciatus
Giinther and 0. tafniolatus ceylonicus Wall.

Oligodon taeniolatus fasciatus (Glinther)

Oligodun fasciatus Giinther, Reptiles of British India, 1864, pp. 208-209, pi. XIX, fig. D
(type locality, Deccan [India]).

Oligodon taeniolatus fasciatus Wall. Oiihidiri Taprobanica or the Snakes of Ceylon, 1921,
pp. 239, 241-242.

Wall, loc. cit., reports this form from Ceylon. I have seen no
specimens from there. It may be recognized by the median series of
rounded spots which pass down the back, some of the anterior ones
being divided. Other lateral marks and small spots are present near
the edge of the ventrals.

Oligodon taeniolatus ceylonicus Wall

PI. XVI, fig. ]

Oligodon taeniolatus (variety) ceylonicus Wall, Oiihidia Tajirolianica or the Snakes of Cey-
Inii, l!i21, p]). 240-247.

A series of specimens, EHT-HMS Nos. 30165-30172, from a point
12-14 miles north of Trincomalee, Ceylon, have been examined. One
s]iecimen, No. 24138, is in the Kansas University Museum of Nat-
ural History. The following table gives scale counts and measure-
ments of the specimens.

Scale Coi^nts and Measurements of O. /. ceylonicus

Sub-

Scale

Total

Tail

Number

Sex

Ventrals

caudals

formula

length

length

30165

$

175

45

19-17-15

414

68

30166

S

175

47

19-17-15

390

63

30167

5

185

42

19-17-15

374

55

30168

i

175

47

19-17-15

384

62

30169

6

171

45

19-17-15

295

47

30170

?

184

44

19-17-15

311

42.5

30171

£

174

51

19-17-15

156

25

30172

$

174

50

19-17-15

146

23

24138 *

(.vs.)

178

36

19-15-15

151

17

Negonibo, Western Province, Cejlon.

The dorsal seven rows are dark brown, traversed by about 43
narrow, darker bands, composed of a series of black brown flecks
on scales, each i)receded by a series of pure white longitudinal flecks.
Sometimes these areas are rather dim, and when broken medially

Taylor: The Snakes of Ceylon

551

Fig. 3. OUgodon taeuiolatus ccylonicm Wall. EHT-HMS No. 30167; 12 mi.

X Tiincomalee, Ceylon, (x 10).

may alternate. Many of the lateral scales may have dark brown,
diagonally placed flecks that tend to form lateral diagonal lines.

A well-defined dark band lies across the neck usually sending for-
ward a median tongue and two lateral prongs which nearly enclose
two elongate light areas on the back of the head; a dark band
crosses the snout and involves the eyes, reaching to the commissure
of the mouth. There is a suggestion of a hair-fine, median line oc-
cupying the median third of the dorsal scale row, but in none is it
continuous. A dim brownish narrow line goes from angle of the

552 The University Science Billetin

mouth to near the vent, lying hugely on the second or third outer
scale rows.

The venter is entirely without dark color. The outer fifth of the
ventrals are creamy Mhite. Along the edge of this color is a trans-
parent line which together with those on other scales form a longi-
tudinal line on each side of the ventrals. Between these lines the
ventrals are flesh color.

The specimens were encountered about rocky outcrops in the
forest. Usually they were found coiled under rocks lying partially
on other rocks. They were very gentle, permitting themselves to be
handled. They did not attempt either to bite or to escape.

The drawing shows the general disposition of head scales.

Oligodon sublineatus Dumeril, Bibron and Dumeril

PL XVII, fig. 1

Oligodon sublirieat^iDi Duiiieiil, Bibron and Dumeril, Erpetologie Generate, vol. 7, pp. .57-58
(type locality, Ceylon); Smith, Fauna of British India, Ceylon and Buriria . . ., Reptilia
and .AmjiliilMa, vol. 3, Serpente.s. 1!»43, pp. 2'27-22S.

A single specimen, U. S. Nat. Mus. No. 56330, ''Ceylon," shows
no divergence of significance from the typical. The jn-esence of the
dorsal narrow transverse paired spots (which frequently alternate),
the three ventral lines of dots (two only on the tail) and the gray-
brown dorsal coloration make this form easily recognized. It has
been remarked that this form shows a marked superficial resem-
blance to Callophis melanurus (Shaw), which also occurs on the is-
land. The following scale characters obtain in this specimen:

Ventrals, 149; subcaudals, 27; anal divided; 7 upper and 7 lower
labials.

The ventral range given by Smith, loc. cit., is 134-161; of sub-
caudals, 23-37.

Oligodon caUvmarius (Linnaeus)

Coluber (-nlainarius Linnaeus, Mus. Ad. Frid., 1754, p. 23, tab. G, fig. 3; and Systema
Naturae, 10th edition, 1754, p. 21(), and 12th edition, p. 375 {fide .\ndersson, Bihang till k.
Svenska vct.-.Akad. Hand., bd. 24, .\fd. IV, No. 6, 1899. pp. 8-9) (type locality, '.America" in
error).

Oligodon tcinpleloiii Giinthtr, Ann. Mag. Nat. Hist., ser. 3, vol. 9, 1862, ji. 57 (t.\pc locality,
Ceylon): and Kept. British India, 1864, p. 209, pi. XIX, fig. c.

Oligodon calatnariu.'s Andersson, loc. cit.; Smith, Fauna British India . . ., Rcjitilia and
Amphibia, vol. 3, Serpentes, 1943, pp. 228-229, fig. 78.

No specimens are in the collection. The species is known from
numerous localities, occurring from the lowlands to an elevation of
3,000 to 4,000 feet.

Taylor: The Snakes of Ceylon

553

■•>

Plate XVII. Fis- 1- Oligodon sublineatus Dumeril. Bibion and Diimeril.
U S.N.M. No. 56300 5 ; "Cevlon"'; total length, 260 mm. Fig. 2. Asjndura
trachyprocta Cope. EHT-HMS No. 30731; "Ceylon"; total length, 316 mm.

554 The University Science Bulletin

Genus Ahaetulla Link

Ahaetulki Link, Beschreil)ung der Natiiialien-Sammlung der UiiiversitiU zu Rostock, vol. 2.
p. 73.

Genotype, fasciata = Coluber ahaetulla.

There has been considerable confusion in the application of the
generic name Ahaetulla/' It is apparent that the name must be ap-
plied to a group of oriental snakes rather than to a group of South
American forms related but not congeneric with them. It is fitting
since the name Ahaetulla presumably is based on a Sinhalese word.

P'our sj^ecies of this genus occur in Ceylon. One is here described
as new. These may be differentiated by the following characters:

Key to Ceyloxfse Species cf Ah.aetili.a

1 . Xo loieal present ; scales in 1.') rows : ventrals 175 oliveri

( )ne or more loreals presert 2

2. Two loreais ; scales at middle of body in 15 rows; ventrals 154-170 bifrenalis

One loreal ; scales in either l.S or 15 rows 13

3. Scale rows 13 ; ventrals 149-104 caudoliiieolatus

Scales in 15 lows ; ventials 103-107 trisiis

Ahaetulla bifrenalis (Boulenger)

Dtndrophis bifrenalis Boulenger, The Fauna of British India, 1890, p. 33S (type locality,
Ceylon).

One specimen (EHT-HMS No. 30745) of this rare species was
captured 13 miles north of Trincomalee.

It may be identified by the character of the double loreal, and the
relatively low ventral count (154-176) and relatively high sub-
caudal count (144-175). The scale formula is 15-15-11. The verte-
brals are strongly enlarged, broader than the outer scale row at
midbody, their posterior margin truncate or concave.

No. 30745. The nasal scale differs from that of other mem-
bers of tiie genus occurring in the same locality. The nostril is near
the middle of a large undivided scale. From the nostril a groove ex-
tends back to the anterior loreal which I suspect may actually be a
very aberrant posterior nasal. The preoculars are 1-1, the post-
oculars 2-2. The temporals are 1 -f 2 + 2 (2 + 1 + 2). Supra-
labials are 9-9, of which the fifth and sixth border the orbit; in-
fralabials are 10-10, five of which border the anterior chinshield.
The eye is large but its diameter is distinctly less than its distance
from the nostril. The ventrals are 168, each with lateral keels.
There are 145 subcaudals, all of which have keels. There is a di-
vided anal.

* An excellent review of this bv Dr. J. Oliver appears in Bull. Anier. Mus. Nat. Hist.,
vol. 02, 1048, ]))). 167-170.

Taylor: The Snakes of Ceylon 555

The top of the head is gray. The side of the head bears a black
stripe that passes through eye to the neck. Anterior labials are pure
cream save on their upper edges. A series of c.ashlike white marks
are present on the first and second scale rows occupying only the
adjacent edges of the scales. A series of dark diagonal marks pass
across the anterior third of the body, none of which are wider than
one scale length.

Aliactidla oliveri sp. nov.

PI. XVIII. fig. 1

Tijpe. EHT-HMS No. 30388 collected 12 miles north of Trin-
comalec, Ceylon, in 1944 by E. H. Taylor.

Diagnosis. Three labials, the fourth, fifth and sixth entering the
eye; loreal absent; nasal divided at least partially; the posterior
part largest and a little lower than anterior; temporals 1 + 1 + 3;
vertebral scales enlarged, smaller than scales of outer row; a lateral
black headstripe continues through eye and to base of tail; a white
line beginning on supralabials continues on the first and second
scale rows, occupying most of first and lower half of second scale
row; this bordered below by a black line beginning on neck and
continuing to base of tail; tail gray without stripes.

Description of species. Rostral large, its width much greater
than its height, part visible above slightly angular posteriorly;
internasals longer than the prefrontals but much narrower; pre-
frontals bordering the second to fourth supralabials; frontal equally
as long as its distance from the tip of snout ; parietals as long as
their distance from internasals; nasal elongate, completely or al-
most completely divided, the posterior portion low^r and larger
than the anterior which contains most of nostril; one large pre-
ocular, touching frontal ; two postoculars, the upper much the
larger; temporals 1 + 1 + 3, the lower of the three almost below
the second temi^oral; sujiralabials 9-9, the fourth to sixth bordering
orbit ; infralabials 10-10, five bordering first chinshields. Ventrals
173; anal divided; subcaudals divided, 134; ventrals keeled and
notched on each side; subcaudals with rounded lateral ridges in lieu
of lateral keels.

Color. Above bronzy gray to middle of third scale row, a cream
white line bordered above and below by deep black passes on two
outer scale rows to tail. Tail uniform bronzy gray. Belly cream
white (in preservation the belly tends to become dark).

Measurements in mm. Total length, 756; tail, 250.

550

The University Science Bulletin

"N*- -rt»Ti.'» .',■-'

Taylor: The Snakes of Ceylon 557

Retiiarks. The type specimen was taken in low forest near the
beach. Its movements were moderately rapid. The species is dedi-
cated to Dr. Jim Oliver who has contributed to the stabilization of
the generic name of this form.

In 1909 * Werner described a Ceylon species lacking a loreal as
Dendrophis effrenis. This species has been referred by most sub-
sequent writers to the synonymy of Ahaetulla candolineolata. Even
Werner does not recognize the form in his "Gattungen und Arten
der Schlangen aus der Familie Colubridae" (Zool, Jahrb., Bd. 57,
1929).

This form belongs to the section of the genus having 15 scale rows
and with three instead of two supralabials entering the eye. Wer-
ner's effrenis belonged to the group having 13 scale rows, with two
labials entering the eye.

Ahaetulla triatis (DaucUn)

Coluber tristis Daudin, Histoiie Naturelle des Reptiles, vol. 6, p. 430 (Based on Russ(;ll.
.\n Account of Indian Serpents, 1796, p. 36, pi. 31).

Chrynopelea boiei A. Smith, Mag. Zool. Bot., 1836, p. 144 (Ceylon).

IJeiidrophit: heleiia Werner, Zool. Anz., 1892, vol. 16, p. 8 (Ceylon).

Dendrclaijhis tristis var. iaprobaneiisis Wall, Ophidia Taprobanica . . ., 1921, p. 221
(Ceylon).

There are seven specimens of this common species in the col-
lections. These are EHT-HMS Nos. 30733-34; 30736-37; 30740-42,
all from 12 miles north of Trincomalee.

A young specimen, U.S.N.M. 12494, differs in color characters from
the others listed in having the anterior vertebral scales each with
a cream spot. Each transverse, somewhat diagonal, row of scales
has the posterior edge black thus forming as many narrow black
lines as there are transverse scale rows; some of the scales have their
lower edges dark also and together these form a series of dim diag-
onal transverse lines but directed obliciuely backwards. This second
series cannot be discerned on the second half of the body.

The two outer scale rows are largely cream but each third scale
of the outer row either black-edged or with the adjacent scales black-
edged so that there appears a series of black-edged yellowish cream
spots. These are present only on the anterior half of body. Whether

* Mitt. Naturh. Miis. Hamburg, vol. 26, 1909. p. 221.

Pl.ate XVIII. Fig. 1. Ahaetulla oliveri sp. nov. EHT-HMS No.
30388(5 ; 12 miles: north of Trincomalee, Eastern Province, Ceylon; total
length, 756 mm. Fig. 2. Chryaopelea taprobanica Smith. EHT-HMS No.
31244,3 ; 12 mile? north of Trincomalee, Eastern Province, Ceylon; total
length, 656 mm.

558 The University Science Bulletin

this is the typical juvenile marking or whether this is an unusual
condition, I cannot say.

The ventrals are 164, the subcaudals 125; the anal is divided.

AhaetuUn caurlolineolata (Giinther)

DendrophiK (■audolincolatut: CUinther, Proc. Zjol. Soc. Ldiulun, iy(i!». p. 507, pi. 11, fig. 1
(type locality, Ceylon).

Dendrophis gregorii Haly, Taprobanian, 1888, vol. 3, p. 51 (Ceylon).

DendrophiK cjjrenis Werner, Mitt. Naturhist. Mus. Hamburg, vol. 20, 1909, p. 221 (Colombo,
Ceylon).

No specimens of this species are in the collections. The species
may be diagnosed by the following characters: scale rows 13-13-9;
eight supralabials, the fourth and fifth borrering eye; anteriorly with
diagonal dark lines; tail with four more or less distinct black longi-
tudinal lines. Ventrals 149-164; subcaudals 119-128; anal divided.

genus chrysopelea Boie

Chrysopelea Boie, in FtTnii^ac, Bull. Sci. Nat., vol. 0, 182(1. 11. '23 7 : and Isis, 1827, p. 520.

Two forms occur in Ceylon. They may be distinguished by the
following key.

Key to Ceylon Species of Chrysopelea

1. Scale preCftUng an:il scale paired; greenish with transverse darker markings form-
ing bands and with a median series of orange spots pre.«ent orituta

Scale preceding anal scale single; light olive brown with aliout (>2 black wavy or

zigzag lines on body taprohaiiiia

- Chrysopelea ornata (Shaw)

Coluber oniatu/' Shaw, Cicneral Zoology, 1802, vol. 3, p. 477 (basetl on Russell's Iml.
Serp. 2, 1801, p. 4, pi. 2; unknown type locality, but jiresumably India).
Chrysopelea ornata Boie, Isis, 1827, p. 546.

No specimens from Ceylon have been examined. The flower-
shaped spots of orange are said to be invariably present on Ceylonese
specimens.

Chrysopelea taprohanicd Smith

PI. XVIII, fig. 2

Chrysopelea taprobanica Smith, Fauna of British India, Ceylon and Burma including the
whole Indo-Chinese Subregion; Reptilia and .\mphil>ia, \-ol. 3. Seriicntes, Dec, 1943, p. 254
(type locality, Kanthali, Ceylon).

Crysopelea ornata (in part) of various authors.

This recently described species is represented in my collection by
a single specimen (EHT-HMS No. 31244) collected 12 miles north
of Trincomalee. It is surjirising that the species was ever confused
with Chrysopelea ornata, which likewise occurs on Ceylon, since the
general appearance is quite different. The ground color of tapro-
banica is light olive brown with a series of 62 transverse wavy or

Taylor: The Snakes of Ceylon 559

zigzag lines crossing the back that often break up low on the sides,
and, at least in the posterior part of the body, they fail to reach to
the ventrals. The greatest width of the black color is usually less
than half a scale in width. These bands continue on the tail, but
become indistinct towards the tip. Thirty or more can be counted.
The ventrals are sharply keeled, notched laterally, and bear a
broken row of black dots touching the keel. These dots are ar-
ranged in groups of three or two on contiguous scales, separated by
a scale without a dot. They continue on the subcaudals but be-
come irregular and are absent posteriorly.

When the outer epidermis is shed the color is light blue-gray. The
head is black crossed by three continuous ivory-colored bands; be-
tween these are other bands made up of ivory flecks or spots and
short lines. The chin and labials are ivory white.

The keels on the scales arc indistinct and if the outer epidermis is
shed can scarcely be discerned. The scale rows are 17-17-13. The
anal is divided and the last ventral is single. The ventrals are 204;
the subcaudals 110 -f (the extreme tip missing).

The species is confined to Ceylon. It is known from the North
Western Province (Kurunegala) and the northern part of Eastern
Province (Kanthalai, and 12 mi. north of Trincomalee). It would
appear to be a lowland form.

Genus Lycodon Boie

Li/roiloii B.iie, in J'Vniisac's Bull. Sci. Nat., vol. 9, 1820, p. 238 (part); Fitzingor, Xciie
Class. Rfpt.. 182(1, pp. 29, 30.
Gcnotyiie, Lycodon auliciis.

Only three species are known to occur in Ceylon.

Key to the Forms of Lycodon in Ceylon

1. Eight siipialabials ; ventrals not aiigulate laterally; ventrals (Ceylon) 154-166;

subcaudals, S.i-.tO; dark brown with whitish crossbars, which widen on sides,
the expansions often enclosing triangular spots; or becoming pigmented; pos-
teriorly bars narrower and closer together; below uniform white strUifiis

Nine supralabials ; ventrals angulate laterally ; ventrals, 172-214; subcaudals, 57-

80 2

2. Brown, lavender, or purplish, with white bands, which widen on the sides; labials

cream without 'spots in their middle; diameter of eye two times in distance

between eye and nostril ; preocular touches frontal aulicus aiiliriis

Brown or lavender with a series of small, whitish, dark bordered medial spots on
anterior two thirds of body ; eye diameter one and one half times in distance
from eye to nostril ; preocular separated from frontal osmanhilli

560 The University Science Bulletin

Lycodon aulicus aulicus (Linnaeus)

PI. XIX. fig. 2

Coluber auHrus Linnaeus, Miistum Adnlphi Fiidcrici regis, vol. 1, 17.54, p. 29, pi. 12, fig. 2
(.America); Systenia Naturae, 10th ed., 1758, p. 222.

Lycodon aulicus Wall, Jouin. Bombay Nat. Hist. See, vol. 15, 1904, p. 706; idein., vol.
18, 1907, p. 112; idem., vol. 19, 1909, p. 87, color pis. 344,619; Smith, The Fauna of British
India, Ceylon and Burma . . ., Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp. 263-
266, fig. 89.

Ophites aulicus Wall. Ophidia Taprobanicn, or the Snakes of Ceylon, 1921, p. 151.

This species is rei)resented by the following specimens: EHT-
HMS Nos. 30708-30713, 12 mi. N. Trincomalee, Ceylon ; U. S. N. M.
No. 31247, "Ceylon."

The series of specimens from near Trincomalee are uniform in
pattern and seemingly all are referrable to L. a. aulicus.

Temporal scales vary somewhat, the formula being 2 -\- 3 -\- 4.

or more frequently assuming a position of -{-4 (3). The

third, fourth and fifth supralabials border the orbit, five infralabials
touch the anterior chinshields, which are somewhat larger than poste-
rior. The scale formula is 19 (anteriorly on neck), 17, 17, 15.

The head is spatulate, flattened, the rostral only slightly visible
above. The nasal is at least partially divided into two nearly equal
parts and the loreal is in contact with the internasal.

The color on the dorsum and sides is lavender or purplish lavender
(tending to turn brown after several years of preservation), crossed
by 13 narrow, cream-white lines one and one half to three scales
wide, that widen on the sides and become confluent with the lighter
color of the venter. The top of the head is deep lavender bordered be-
hind by a cream transverse band that is divided by a narrow tongue
of lavender. The lips are cream except on the uj^per edges. Older
specimens develop pigment on the scales of the white bands and the
white may form only a reticulum; the borders of the lavender
areas may be darkened and irregular.

Wall, loc. cit., has called attention to the variation occurring in
Indian and Ceylon specimens. It is surprising that the series here
shows very little variation in color save that normally expected with
age.

Pl.ate XIX. Fi^;. 1. Sibynuphi.s subpunclatus (Diimoril. Bibron ami
Dumeril) KHT-HMS No. 31252 9; 12 miles north of Trincomalee,
Eastern Province, Ceylon; total len^ith, 355 mm. Fifi. 2. Lycodon
aulicus aulicus (Linnaeus). EHT-HMS No. 30712 9 ; 12 miles north of
Trincomalee, Eastern Province, Ceylon; total length, 378 mm.

Taylor: The Snakes of Ceylon

561

35—90

562 The University Science Bulletin

Table of Data on Lycodon auHciis auHcus (Linnaeus)

Sub-

Supra -

Infra -

Number

Sex

Ventrals

caudfils

labials

labials

Oculars

30708

S

185

67

9-9

11- 9

1. 2

30709

S

184

65

9-9

11-11

1, 2

30710

S

180

67

9-9

11-11

1, 2

30711

9

201

63

9-9

10-11

1, 2

30712

9

195

56

9-9

11-10

1. 2

30713

S

181

67

9-9

11-10

1, 2

31247

s

185

69

9-9

11-11

1, (2-3)

Lycodon striatiis (Shaw)

Coluber striafus Shaw, General Zoology, vol. 3, 1802, p. 527 (Based on Russell's, An account
of Indian Serpents . . ., pi. 16).

Lycodon striatiis Smith, The Fauna of British India . . ., Reptilia and Ainphil)ia, vol.
3, Serpentes, 1943, pp. 20] -202. Ceylon (etc.).

No specimens of this species in the collection.

It may be distinguished by the presence of only eight supralabials,
and in having the anterior part of nasal larger than posterior. The
rounder rather than angulate condition of the ventral scales will
separate this species from Lycodon aulicus. The ventrals are 185,
subcaudals 69 -|- 1; the supralabials 9-9, the infralabials 11-11.

There is a single preocular but on the left side there are three
postoculars instead of two, the usual number. The arrangement of

the temporals is- — --^ -|- 4. The third, fourth and fifth supra-

labials bound the orbit.

Lycodon osmanhilli sp. nov.

PI. XX

Tyve. K. U. M. N. H. No. 24141, Colombo, Ceylon; W. C. Osman
Hill, Coll., paratype U. S. N. M. No. 19215, "Ceylon."

Diagnosis. Diameter of eye (2.9 mm.) contained in distance be-
tween eye and nostril (4.0 mm.) 1.38 times; length of frontal equal
to distance between frontal and rostral; tail (101 mm.) in total
length (522 mm.) 5.38 times. Ventrals angulate; nine supralabials,
each with a medial black spot; a series of small, dark-bordered
cream spots on anterior two thirds of body; preocular separated
from frontal.

Description of type. Rostral forming a sharp transverse ridge, its
width twice as great as its height, the upper part bent at nearly a
right angle, rather broadly visible from above; internasals a third
longer than wide forming an angular suture with the rostral; pre-
frontals approximately one third longer than broad, their area more
than three times that of the internasals; frontal shield-shaped

Taylor: The Snakes of Ceylon

563

Plate XX. Lycodon OKmanhiUi sp. nov. K.U. M.N.H. No. 24141; Colombo.

Ceylon; total length, 522 mm.

564 The University Science Bulletin

nearly a third longer than wide, its length equal to its distance from
rostral, and about one fourth shorter than a parietal ; latter scales
much narrowed posteriorly. Nasal divided, the nostril pointing up-
ward and somewhat outward; anterior part of nasal largest; loreal
elongate, more than twice as long as high; not in contact with the
internasal; one preocular as high as wide, separated from frontal;
two postoculars ; temporals 2 + 3-1-4, those bordering the parietals
not especially enlarged; supralabials 9-9, the third, fourth and fifth
bordering the eye; infralabials 10-11, five bordering the first pair
of chinshields, wiiich are distinctly larger than second pair; neither
the snout nor lower jaw constricted immediately in front of eye.

Across back of head 23 scale rows; on neck and body 17; in front
of vent" 15. Ventrals angulate, 187; anal divided; subcaudals 68.
Scales smooth ; single apical pits are discernible if the outer epider-
mis is not shed.

Color. Tiie general color brown in alcohol, the top of head being
uniform lavender brown, bordered along the upper edge of labials
by a slightly darker line. All labials cream, with a darker upper
edge and large dark medial spots; infralabials each with a dim
darker spot; venter immaculate. Back brown with 19 cream spots
covering part of three or four scales on middorsal line, each edged
with dark brown; many lateral scales with very narrow white edges;
latter third of body and tail lacking spots. Most of the dorsal and
lateral scales are of a darker brown on their edges than in the
middle.

Measurements in mm. Total length, 522; snout to vent, 421; tail,
101; greatest width of head, 10.6; greatest length of head (from jaw-
angle) 17.8.

Remarks. The paratype, U. S. N. M. No. 19215, is a young, per-
haps somewhat faded specimen. However the general color pattern
of the young is identical to that which obtains in the adult. There
are 21 median white or cream spots, covering parts of four scales
and completely or partially surrounded by dark brown. The dark
labial spots are distinct.

The eye (as is usual in many snakes) is proportionately larger
than in the adult and its diameter is only minutely less than distance
between eye and nostril. The preocular is separated from the
frontal, and other head scales have the general shape and propor-
tions as those of the type.

The following scale characters are present: supralabials, 9-9; in-
fralabials 11-11, five touching the first chinshields; one loreal; one

Taylor: The Snakes of Ceylon 565

preocular; 2-3 postoculars; scale formula, 19-17-17-15. Ventrals,
186; subcaiidals (divided) 70 (the terminal subcaudal scale eciual
to the length of the preceding five pairs of subcaudals) ; anal divided.

Co77iparisons. The species here treated as Lijcodon milieus aidicvs
although having a somewhat similar scale formula differs in pat-
tern, in the size of eye, which is smaller, in the constriction of the
snout in front of or at eye. The eye is contained in the nostril to
eye distance more than twice, the eye being smaller and the distance
greater than in osmanhilli. When two specimens of near equal
length arc compared, Lycodon aulicus aidicus is more robust, witli
a broader and a longer head. The supralabials and infralabials
lack the median dark spots.

Two specimens of mdicus, one a little longer and one a little
shorter, are compared by a series of measurements.

Comparative Measurements of Lycodon oamanliiUi and L. aulii'iis nulicns

Head

E>e to

Tail

Totul

Hesd

width

Eye

.snimt

Eye to

in total

Number

length

Tail

length

greatest

diameter

tip

nostril

length

30710

545

98

21.1

13.2

2.9

7.1

5.45

5.56

24141

522

99

17.8

10.6

2.9

5.8

4

5.38

30713

498

95

19.2

11.3

2.2

6.3

4.5

5.24

Nos. 30710, 30713 — L. auliots nnlicits from 12 mi. X. Tiincomaloo.
No. 24141— Type of os,)HnihiUi.

In Lycodon aulicus aidicus the preocular is much higlier than long
and is in contact with the frontal. The temporals bordering the
parietals are enlarged, there being only three on each side (four in
osmanhilli). The loreal is deeper, its width minutely less than
twice in length. The length of the frontal equals distance from
the frontal to the middle of the internasals. There is no significant
difference between the counts of the ventrals and subcaudals in the
two forms. The counts of the ventrals in males are 181-185 (186-
187) ; subcaudals 65-69 (68-70) ; in the females, 195-201; subcaudals
56-63 (counts for osmanhilli in parenthesis).

The species is named for Dr. W. C. Osman-Hill of Edinburgh,
Scotland, who has collected widely in Ceylon.

Genus Cercaspls Wagler

Cercaspis Wagler, Natlirliches System der .\m]iliiliien. ISSO. p. !(1.
Genotype, Hurria carinata Kuhl.

Cercaspis carinatus (Kuhl)

Hurria carinata Kuhl, Beitr. Zool. Vergl. Anat., 1820, p. 95 (tjpe locality unknown).

Cercaspis carinattis Giinther, Reptiles of British India, 1864, p. 324; Wall, Spolia Zeylanica,
vol. U, 1921, pp. 399, 404; and vol. 13, 1924, p. 77; Ophidia Taprobanica, or the Snakes of
Ceylon, 1921, pp. 162-16.';; Joum. Bombay Nat. Hist. See, vol. 29, 1923, p. 614; Smith,

566 The University Science Bulletin

Fauna of British India . . ., Reptilia and Amphibia, vol. 3, Serpentes, 1943. p. 268-269,
fig. 90 (vertebrae).

Lycodon carinatus Boulenger, Fauna British India, 1890, p. 297; and Catalogue of the
Snakes in the British Museum, vol. 1, 1893, p. 358.

This species resembles closely in external characters certain spe-
cies of Lycodon but differs in an extraordinary manner in having
the very greatly broadened prezygapophyses on the vertebrae.
Smith {loc. cit., footnote p. 267) states that this same condition is
present "also in S. American Xenopholis."

No specimen in the collection.

Genus Dryocalamus Giinther

Dryocatuinuii Giinther, Catalogue of the Colubrine Snakes in the Collection of the British
Museimi, 1858, p. 121.

Genotype, Dryocalamus tristriyatus.

Two species occur in Ceylon, neither of which is endemic.
Key to the Ceylon Forms of Dryocalamus

Scales in 13 rows; one or two preoculars; nasal undivided or partially divided. . . . nympha
Scales in 15 rows; one preocular; nasal divided more or less completely gracilis

Dryocnlanius nympha (Daudin)

Coluber nympha Daudin, Histoire Naturelle des Reptiles, 1803, vol. G, p. 244, pi. 75.
Dryocalamus nymplia Boulenger, Catalogue of the Snakes in the British Museum, vol. 1,
1893, p. 370.

No specimens in the collections.

These smooth-scaled snakes reach a length of about half a meter.
The nasal is not or only partially divided below the nostril. The
scale formula is 13-13-13. The head is flattened and distinctly
wider than the body. Ventrals 200-236; subcaudals 65-88.

Dryocalmnus gracilis (Giinther)

Adoiitoinits yracilis Ginther, The Reptiles of British India, 1804, p. 234 (type locality,
Anamally Hills).

Dryocalamus gracilis Boidonger, Catalogue of the .'>nakes in the British Mu.seuni, vol. 1,
1893, p. 371.

No specimens in the collection.

In this species the nasal shield is more or less completely divided.
The loreal is in broad contact with the orbit and has usually a small
preocular above it bordering orbit. The scale formula is 15-15-15.
Ventrals l!)i)-243; subcaudals, 75-87.

Genus Natrix Laurenti

Katrix Laurenti, Specimen Medicuni, exhibens Synopsin Reiitilium emciKhitun]. 17(18, p. 73.
CJenofype, Natrix vulgaris = Coluber natrix Linnaeus.

Three forms are known to occur in Ceylon. These may be dif-
ferentiated by the following key.

Taylor: The Snakes of Ceylon 567

Key to Ceylon Species of Natrix

1. Posterior maxillary teeth enlarging gradually; no dorsolateral buff stripes; two

diagonal lines below and behind eye 2

The last two or three posterior maxillary teeth abruptly enlarged ; usually buff
dorsolateral stripes present, more distinct posteriorly ; no diagonal lines below
and behind eye stolata

2. Anterior half of body pale olive or reddish, with two series of more or less distinct

alternating large roundish or rhomboidal, dark olive or brown, black-edged
spots, which are partly confluent on the vertebral line (sometimes forming a
sinuous band); posterior part of body of uniform color or with longitudinal
rows of dark spots; 2 diagonal lines below and behind eye; ventrals, 131-146;

subcaudals, 83-90 asperrimus

Anterior half of body with a dim black-edged nuchal band, followed by a median
darker spot with a lighter center ; behind this six series of irregular darker
spots arranged in longitudinal lines, the individual spots often dim and often
confluent; series of narrow vertical light spots sometimes confluent on the
median line. Ventrals, 131-140 ; subcaudals, 78-90 piscator piscafor

Natrix asperrimus (Boulenger)

PI. XXI, fig. 1

Tropidonotus asperrimus Boulenger, Ann. Mag. Nat. Hist., .ser. 6, vol. 7, 1891, p. 281; and
Catalogue of the Snakes in the British Museum, vol. 1, 1893, p. 232, pi. 15, fig. 2 (type
locality, Ceylon).

Natrix piscator asperrimus Smith, The Fauna of British India . . ., Reptilia and
Amphibia, vol. 3, Serpentes, 1943, p. 290.

Two specimens (U. S. N. M. Nos. 120329-30) are present in the
collections. The form may be recognized by the characters pre-
sented in the Key.

Unless it can be shown that in Ceylon piscator and asperrimus
have ranges distinct from each other and show intergradation, I
regard it as necessary to consider them distinct species, as proposed
by Boulenger.

The head of No. 120329 is dark blackish brown to black, the
color continuing about nine scale lengths behind the parietals; the
whitish nuchal band is three scales wide on the middorsal line, then
widens on the sides and becomes confluent with the ventral colora-
tion. A pair of diagonal lines are under eye, enclosing a white area,
the upper being continued as a border on the dark spot of the
head, the other is traceable across the jaw. A series of dashlike
black marks are present on the edges of the ventrals and the first
scale row. The series of dorsal spots are separated by transverse
light bars for nearly half the length of the body. Posteriorly the
large spots gradually disappear, and a few narrow indistinct dark
markings can be discerned together with a series of very small white
spots present on each side to the tail. The subcaudals are bordered
by a black line. Ventrals 133 ; subcaudals 72.

No. 120330 has become nearly unicolor but some trace of the
markings are visible when submerged in water.

568

The University Science Bulletin

'^s^ii^sX.^K*"

Taylor: The Snakes of Ceylon 569

Natrix piscator (Schneider)

PI. XV, fig. 2

Ilyilrtis pmcator Schneider, Histoiia Aniphibioruni natuialis et literaria, vol. 1, 17'J9, jj. 247
(Based on Russell's, An Account of Indian Serpents, pi. 33).

Natrix piscator piscator Smith, Rec. Indian Mus., vol. 42, 1940, p. 483 ; and The Fauna of
Biitish India . . ., Reptilin and Amjihihia, vol. 3, Serpentes, 1944, p. 29.').

Smith, in his recent Fauna of British India, does not consider this
species or subspecies as occurring in Ceylon. A series of specimens.
Nos. 20724-20728, 30730, which I collected 12 miles north of Trin-
comalee, are referred to Natrix pi'scatof and seemingly are more
closely related to the typical form than to any other as regards the
pattern. One specimen, No. 30729, was collected between Anurada-
pura and Colombo, about 50 mi. S of the former city.

T.^BLE OF Data on Natrix ])/.'<c(dar

Suh-

Suiira -

Infra -

Pre-

Post-

Number

Sex

Vontral.s

oauduls

labials

labials

iicular

(icnlars

30724

9

139

85

9-9

10-10

1-1

3-3

30725

9

139

53 +

9-10

11-10

1—1

3-3

30726

yg-

131

47 +

9-9

10-10

1-1

3-3

30727

9

138

87

9-9

11-10

1-1

3-3

30728

9

140

S3

9-9

10-10

1-1

3-3

30729

S

134

90

9-9

11-10

1-1

3-3

30730

9

136

78

9-10

10-10

1-1

3-3

The pattern of dorsal spots is distinct in certain specimens, less so
in others. In all, the spots are more or less contiguous, and not of
solid color. The black usually outlines three or four scales whose
centers are much lighter. The small narrow vertical white spots
(never as wide as one scale length) are present on the sides of all
the specimens, usually as far as the base of the tail. They can be
discerned even when the pattern of dark spots is indistinct. In one
specimen the white spots tend to meet on the middorsal line, and
they are least conspicuous in the specimen from the western side of
the island south of Anuradapura.

Plate XXI. Fig. 1. Natrix aspenimus (Bou\enger). U. S. N. M. No. 120329;
NancUma Estate, Kandv District, Central Province, Cevlon; total length, 252
mm. Fig. 2. Natrix stolata stolata? EHT-HMS No. 30706 9 ; 12 miles north
of Trincomalee, Eastern Province, Cevlon; total length, 415 mm. (tail light
below). Fig. 3. Natrix stolata stolata (Linnaeu.s). EHT-HMS No. 30705;
Kandy, Central Province, Ceylon; total length, 440 mm.

570 The University Science Bulletin

Natrix stolata stolata (Linnaeus)

PI. XXI, figs. 2 and 3

Coluber stolalus Linnaeus, Systema Naturae, 10th ed., 175S, p. 219 (type locality, Asia).
Natrix stolata Smith, The Fauna of British India . . ., Reptilia and Amphibia, vol. 3,
Serpentes, 1943, pp. 303-305.

Three specimens in the EHT-HMS collection from Ceylon: No.
30705 from Kandy ; Nos. 30706-07 from 12 mi. N of Trincomalee.

The specimen from Kandy, Ceylon, is gray (brownish gray before
loss of the epidermis) , the edges of some of the scales and the inter-
vening skin between some of the scales whitish. Near the head a
lighter line begins on each side covering approximately half of the
fifth and seventh scale rows and all of the sixth row on the anterior
two thirds of the body. Posteriorly the stripe is on the fourth, fifth
and sixth rows to the vent. The stripe continues to the tip of the
tail. Anteriorly the scales of the stripe are dark edged and a chain-
like pattern is in evidence on the white stripe; posteriorly the dark
markings are lacking on the stripe. A black dot is present on the
outer edge of the ventrals on the anterior half of the body. Between
the stripes there is a series of transverse dark markings having ir-
regular edges (these are continued below the dorsolateral line). The
ventrals are $ 143, the subcaudals 77 with possibly a few more
missing from the tip.

The specimens from Trincomalee differ from the specimen de-
scribed in having the dorsolateral light line indistinct throughout the
anterior part of body but beginning one or two headlengths behind
the head is a series of buff spots slightly darker edged which continue
back two thirds the length of the body. Posteriorly the stripe is more
distinct but considerable pigment is scattered on the scales of the
stripe and the lateral spots are more or less indicated. Darker
markings between the stripes can be discerned with difficulty. A
second row of lighter spots can be discerned on the outer scale row.
These also are bordered with darker color. The dark marks on
outer edge of the ventrals appear on only a few scales. The mark-
ings on the sides of the head consist of a vertical light line in front of
and behind eye. The labials are cream with dark linear spots on
the first six that tend to border the sutures.

The scales of Nos. 30706 and 30707 are respectively: ventrals, 125,
126; subcaudals, 66, 69.

The specimens are insufficient in number to determine whether
these differences of color are constant and whether there is as large
a constant difference in the ventrals and subcaudals as is here indi-
cated (maximum 25 ventrals and subcaudals) . Usually females have

Taylor: The Snakes of Ceylon 571

a larger number of ventrals and the count of males of the lowland
form will probably be even less than the low figures here given for
the females.

Genus Balanophis Smith

Balanophis Smith, Proc. Zool. Soc. London, 1938, p. 583.
Genotype, Tropidonotus ceylonensis Gi'inther.

One species known, endemic in Ceylon.

Balanophis ceylonensis (Giinther)

Tropidonotus chrysargus ceylonensis Giinther, Catalogue of the Colubrine Snakes in the Col-
lection of the British Museum, 1858, p. 71 (type locality, Ceylon).

Balanophis ceylonensis Smith, Proc. Zool. Soc. London, 1938, p. 538; and The Fauna of
British India . . ., Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp. 310-311, fig. 98,
A and B.

This rare snake has a series of 24-26 maxillary teeth followed
by two enlarged grooved fangs. It may be diagnosed by the fol-
lowing characters. Scales in 19 rows, all except outer row with keels;
two preoculars, three postoculars ; eight supralabials, four infra-
labials touching first pair of chinshields; ventrals, 131-141; sub-
caudals, 40-54; anal divided. Olive brown above, with more or less
distinct reticulated, black cross bars, enclosing a dorsolateral series
of large yellow or reddish black-edged spots. A dark brown stripe
from behind eye on neck ; skin between scales scarlet, the color evi-
dent when the snake inflates its body.

Genus Macroplsthodon Boulenger

Macropisthodon Boulenger, Catalogue of the Snakes of the British Museum (Natural His-
tory), vol. 1, 1893, p. 265.
CJenotype, flaviceps.

Only one of the four species of the genus is found in Ceylon. It
is also distributed widely in India.

Macropisthodon plumbicolor (Cantor)

PI. XIII, fig. 2

Tropidonotus phimbicolor Cantor, Proc. Zool. Soc. London, 1839, p. 54 (type locality, Mahva
[Saugor], Central India).

Macropisthodon plumbicolor Boulenger, Catalogue of the Snakes in the British Museum
(Natural History), vol. 1, 1893, pp. 207-268; Smith, Fauna of British India, Ceylon and
Burma, including the whole Indo-Chinese subregion; Reptilia and Amphibia, vol. 3, Serpentes,
pp. 314-316, fig. 99, A and B.

I obtained a single specimen in second growth forest 12 miles north
of Trincomalee (EHT-HMS No. 30681). The scale formula is 27,
(head), 23, 23, 17. There are 151 ventrals and 34 subcaudals, and
the anal is divided. There is only a single preocular, the loreal en-
tering the eye. There are three postoculars present and the temporals

572 The University Science Billetin

are 2 + 3. The fangs of this form are very greatly enlarged and are
directed backwards in the mouth.

The color in life was nearly uniform dull green above with some
suggestion of darker bands on the anterior part of the body. On
distending the skin these are well defined. Light pinkish areas are
discernible on the sides on the anterior half of the body. The two
outer scale rows are lighter than the others. The belly is uniform
light cream with a minute dusting of darker pigment on the outer
part of the ventrals. The specimen is a female.

The range of ventrals and caudals for Ceylon specimens given
by Smith (loc. cit.) is: ventrals, 153-154; subcaudals j 40-45;
2 37-47 based on 5 Ceylon specimens. The young are strongly
marked with a V-shaped spot on the neck, followed by an area of
bright orange. Numerous black bars run across the body and tail.
The species is rare in the plains regions, but is encountered fre-
quently in the mountains. Its range extends to an elevation of
7,000 feet.

Genus Atretium Cope

Atretium Cope, Proo. Acad. Nat. Sri. Philadolphia, 18(J1, p. 209.
Genotype, schist osiim.

Atretivm schistosum (DaudinI

Coluber srhistosus Daudin, Histoire Naturelle des Reptiles, \(]1. 7, 1S03. p. 132 (Based on
Kussell's An Account of Indian Serpentes. \ol. 2 pi. 4).

Atretium scliistosus Giinthcr, Reptiles of British India, 18()4, p. 273.

This species may be recognized easily by the fused internasals.
valvular nostrils directed more or less upward, scales keeled without
apical pits, one preocular, two or three postoculars, eight or nine
supralabials, the scale formula 19-19-17 all more or less distinctly
keeled, strongly so on ])osterior part of body. Ventrals 129-160;
subcaudals 53-85. Anal divided.

No specimens in the collection.

Genus Aspidura Wagler

Aspldiira Wagler, Natiirliches System der Anipliiliien. 183(1, jip. 132, 191.
(Genotype, hr(i( liyorros Boie.)

This genus has long been regarded as confined to Ceylon. How-
ever, Laidlaw* rci)orts the species in the Maldive Archipelago. It
is not impossible, if the data are correct, that the species has been
introduced from Ceylon, since it is the snake species most common
in that island and in places it is reported as "exceedingly common."

* Laidlow, in Gardiners, The Fauna and Geography of the Maldive and Laccadive Archi-
pelagoes, vol. 1, pt. 2, 1902, Amphibia and Reptilia, pp. 119-122.

Taylor: The Snakes of Ceylon 573

KiiY TO THE Species of Aspidura

1. Scales in 15 rows ; a preocular present t rarhyprocta

Scales in ] 7 rows 2

2. Two postoculars touching the parietal 3

Upper postocular touching the parietal; a pieocuhu ; snout pointed; suhcaudals

single (lucntheri

3. A preocular present; supraocular nioic than half length of frontal; snout rounrled.

hrn(li!/<irrluin

Xo lucocular ■•

i. Supraocular not half Icnuth ui frontal; .-nout rounded; suhcaudals uinli\ided

(usually) i'op"

Supraoculars more than half length of frontal; snout pointed; suhcaudals paired,

(Inimniofidliayi

Aspiclurn drummondhayi Boulenger

Aspidura drummond-hai/t Houlenger, Spolia Zeylanica, vol. 2, 1904, p. 95, pi. ■ (type

locality, Balangoda district, Ceylon) ; Wall, Ophidia Taprohanica or the Snakes of Ceylon.
1921, pp. 213-214; Journ. Bombay Nat. Hist. Soc, vol. 29, 1923, p. 611; Smith, The Fauna
of British India, Ceylon and Burma including the Whole of the Indo-Chinese subregion ; Reptilia
and Amphibia, vol. 3, Serpentes, Dec, 1943, p. 338.

No specimens examined.

Aspidura trachyprocta Cope

PI. XMI. tig. 2

Aspidura trarli yprocta Cope, Proc. .\cad. Nat. Sci. Philadelphia, 1860, pii. 75-7(1 (type
locality, Ceylon); Giinther. Reptiles of British India, 1864, p. 203, pi. 18, figs. F, F' ;
Boulenger, Fauna of British India including Ceylon and Burma, Reptilia and Batrachia, 1890,
p. 290 ; and Catalogue of the Snakes of the British Museum (Natural History), vol. 1, 1893,
p. 313; Fletcher, Spolia Zeylanica, vol. 5, 1908, p. 98; Wall, Ophidia Taprohanica or the
Snakes of Ceylon, Colombo, 1921. pp. 209-213; Smith, Fauna of British India, Ceylon and
Burma, including the whole Indo-Chinese subregion; Reptilia and Amphibia, vol. 3, Serpentes,
p. 337, fig. 106.

Five specimens, U. S. Nat. Miis. Nos. 19214 Juv. and 56150 Q
"Ceylon"; EHT-HMS Nos. 31245 ^^ and 31246 $ "Ceylon"; and
K. U. M. N. H. No. 24139 j Nuwara Eliya, Ceylon, are at hand, all
falling within the limits of variation of the species.

The type specimen according to Cope's description has two un-
usual characters: a central plate separating the second chinshields,
and a small central postanal plate. These characters at least are
lacking in the specimens at hand. The characters of the scales on
the sides of the anal plate, and the postanal plates do vary however.
In one. No. 31245, there is a small pair of scales inserted between
the outer part of the anal plate and the ventral scale preceding; on
each side of the anus, and partly concealed by the anal plate are
four scales, increasing in size posteriorly. However the pairs fail
to meet. The first postanal is large and single. Towards the end
of the tail there are two divided subcaudals. In the female, No.
31246, the four scales border the side of the vent, but the last pair
is in contact; the intercalated scale between the anal and the pre-
ceding ventral is present on one side only. In No. 56150 there are
no intercalated scales, and but three scales -border the vent, the last

574 The University Science Bulletin

pair not in contact beliind the anus. The first two subcaudals are
divided. The females lack the clawlike keels or tubercles on the
lateral scales preceding and following the vent.

The ventral and subcaudal counts for Nos. 19214 Juv., 31 245 $ ,
31246 5 , and 56150 ? , respectively, are: 142-23; 141-20; 135-13;
143-16.

These small snakes may be easily recognized by the sharp snout,
with a single internasal. The rostral is very small and is visible
above only as a point. The nostril is pierced between two nasals
and the first labial. The scale formula is 15-15-15.

No. 31246 ^ has a single median discontinuous line and a median
ventral irregular series of dark brown spots.

Nos. 56156 ^ and 31245 ^^ have two or three or more irregular
series of small blackish dorsal spots. A brownish line passes back
from eye on to the neck. The venter of the second specimen has
considerable pigment scattered across each scale and not forming
discrete spots.

No. 24139 is dark, nearly uniform bluish black with a row of
eight spots on the third scale row; the belly is light (red or yellow
in life ?) with very numerous quadrangular black spots varying in
size.

Whether the black specimen is racially distinct from the remain-
der listed, I do not know. The cephalic scale pattern is faithfully
duplicated in all. The rostral from Nuwara Eliya is truncate above
while in the others it is pointed and touching the fused internasal
at a single point. Unfortunately the provenance of the other speci-
mens are unknown.

Aspidura brachyorrhus (Boie)

Scytale hrachyorrliof: Boie, Isis von Oken, 1827, p. 517.

Aspidura brachyorrhos Boulenger, Fauna British India including Ceylon and Burma, Reptilia
and Batrachia, London, 1890, p. 288; Catalogue of the Snakes in the British Museum (Natural
History), vol. 1, 1893, p. 311; Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921, pp.
204-207, fig. 43; and Journ. Bombay Nat. Hist. Soc, vol. 29, 1923, p. 611.

Calanmria scytale Schlegel, Essai sur la Physionomie des Serpens, vol. 2, 1837, p. 42 (based
on Boie's Scytale brachyorrhos).

No specimens examined.

Aspidura guentheri Ferguson

Anpidura fiueitthcri Ferguson, Proc. Zool. Soc. London, 187(>, p. 819 (tyiie locality. Coast
of the West Province, Ceylon) ; Boulenger, Faima British India, including Ceylon and Burma
Reptilia and Batrachia, 1890, p. 290; and Catalogue of the Snakes in the British Museum,
vol. 1, 1893, p. 312, pi. 20, fig. 3; Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921,
p. 208-209; and .lourn. Bombay Nat. Hist. Soc, vol. 29, 1923. p. 611; Smith. Fauna British
India, Ceylon and Burma including the whole of the Indo-Chinese subregion ; Reptilia and
.Amphibia, vol. 3, Serpentes, Dec, 1943, p. 338.

No specimens at hand.

Taylor: The Snakes of Ceylon 575

Aspidiira copii Giinther

Aspidiira copii Giinther, Reptiles of British India, 1864, p. 203, pi. 18. fig. E (type locality
.Ceylon); Boulenger, Fauna British India including Ceylon and Burma, Reptilia and Batrachia,
1890, p. 289; and Catalogue of the Snakes L^f the British Museum (Natural History), vol. 1,
1803, pp. 311-312; Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921, pp. 207-208;
and Journ. Bombay Nat. Hist. Soc, 29, 1923, p. 611; Smith, Fauna British India . . .,
Reptilia and Amphibia, vol. 3, Serpentes, 1943. pp. 336-337.

No specimens at hanci.

Genus Haplocercus Gunther

Haplorercus Giinther, Catalogue of the Colubrine Snakes in the Collection of the British
Museum. 1858. p. 290.
Genotype, ceylonensis.

A single recognized form is endemic in Ceylon. It is said to be
common in certain highland districts.

Haplocercvs ceylonensis Gunther

Haplocercus ceylonensis Giinther. Catalogue of the Colubrine Snakes in the Collection of
the British Museum, 18.58, p. 14 (type locality, Ceylon); and Reptiles of British India, 1864,
p. 204, pi. XVIII, fig. G; Boulenger, Fauna of British India including Ceylon and Burma;
Reptilia and Batrachia, 1890, p. 291 ; and Catalogue of the Snakes in the British Museum
(Natural History), vol. 1. 1893, pp. 309-310: Wall, .lourn. Bombay Nat. Hist. Soc, vol. 29,
1923, p. 610, and Ophidia Taprobanica or the Snakes of Ceylon, 1921, p. 143-146, fig. 33;
Smith, Fauna of British India, Ceylon and Burma mcluding the whole of the Indo-Chinese
subregion; Reptilia and Amphibia, vol. 3, Serpentes, Dec. 1943. pp. 340-34^1, fig. 109.

Aspidura carinata Jan, Arch. Zool., 1862, 2, p. 30; and Elenco Sistematico Degli Ofidi,
1863, p. 35 ; and Iconographie Generale des Ophidiens, livr. 13, 1865, pi. I. fig. 5 (type
locality, Ceylon).

Aspidura ceylonensis Theobald, Descriptive Catalogue of the Reptiles of British India, 1876,
p. 143.

A single Ceylonese specimen, U. S. Nat. Mus. No. 5896 lacks exact
locality data. In color and pattern it agrees with Glinther's ex-
cellent figm-e {loc. cit.), and falls within the known limits of vari-
ation of the species. The ventrals are 204, the anal single, the sub-
caudals 44.

The relationship of this genus is apparently with Aspidura. How-
ever much the resemblance with that genus, the differences are such
as to warrant the separation into a separate genus. The variation
in the ventral count is high — 277 to 307 ; while the variation in the
subcaudals is 37-56. The females have a ventral range 192-207;
subcaudals, 37-50; males, ventrals 177-180; subcaudals 42-56. The
greatest variation in number of scales is between a male* with a
ventral-subcaudal count of 219 scales and a female wath 257, repre-
senting a total difference of 38. It is possible that so large a dif-
ference is indicative of racial differentiation.

* Boulenger Catalogue, loc. cit., p. 310.

576 The University Science Bulletin

Genus Boiga Fitzinger

Boiga Fitzinger, Neiie Class. Kept., 1826, pp. 29, 30, CO.
Genotype, Coluber irreoulcwis Meireni.

Five forms of the genus are known to occur in Ceylon.
Kky to the Ceylonese Species of Boiga

1. Three preoculars ; scale rows 19 at miclbod.y reducing to 15 near vent; 4th and

5th labials enter orbit ; length, 2 feet (?) '. barneii

One preocular ; scales 19-27 at midbody ; 3d, 4th and 5th labial,'^ enter orbits.... 2

2. Scales in 19-21 rows ; head color variable 3

Scales in 25-27 rows at midbody; a medial and two lateral stiipes on the head;

length, 7 ft forsteni

3. Scales in 21 rows reducing to 15 posteriorly; \ertebrals enlarged but slightly,

trigonatus
Scales in 19 rows reducing to 15 (Ki) iiosleriuily ; vertebials large, nearly as broad

as long 4

4. Head powdered brown without spols; total ventrals and subcaudals, 362-390;

length, 3.5-4 tt beddomei

Head with dark markings: total ventrals and sulicaudals. 311-342; length about

4 ft ceylonensis

Boiga barnesi (Gunther)

Dipsas barnesi. Giintlier, Proc. Zool. Soc. London, 1869, p. 506, pi. xl, fig. 2 (type locality,
Ceylon).

Boiga barnesi Smith, Fauna of British India, Ceylon and Burma . . .; Reptilia and
Amphibia, vol. 3. Serpentes, 1943, pp. 354-355.

A rare species in Ceylon. Smith states that it i< known only
from two specimens. Ventrals 208-220; caudals, 98-100.

Boiga trigonata (Schneider)

Coluber trigotwtus Schneider, in Bechstein, transl. Lacepede, vol. 4, 1802, p. 356, pi. xl.
fig. 1 (type locality, Vizagapatam, India).

Boiga trigonata Nikolski, Faune de la Russie, 1916, p. 187, pi. VI (trigonatiim); Smith.
Fauna of British India . . .; Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp. 349, 350,
figs. 111. a-d.

Kncwn only I'roin Vvd pr(.^in(■e in Ceylon.

Boiga beddomei (Wall)

PI. XXH. fig. 1

Dipsadomorplnis bedduniei Wall, Records of the Indian Museum, 1909, p. 152; and Ophidia
Taprobanica or the Snakes of Ceylon. 1921, pp. 282-283 (type locality, Ceylon).

Boiga ceylonensis Smith, The Fauna of British India, Ceylon and Burma . . . ; Reptilia
and Amphibia, vol. 3, Serpentes, 1943, pp. 351-353.

Two speeinien.s (EHT-HMS Nos. 30674-75) weie captured 12
miles north of Trincomalee, Ceylon. The first was collected from
under the bark on a standing, dead tree at an elevation of eight feet

Pl.\te XXII. Fis. 1. Boiga hcddomci Wall. EHT-HMS No. 30675 $ ;
12 inilos north of Trinconialeo, Eastern Provinco, Ceylon; total length,
1,190 mm. Fig. 2. Boign fors-tcni Dunieril, Bibron and Dumeril. EHT-
HMS X^o. 30676 $ ; 12 niile.-^ north of Trincomalee, Eastern Province,
Ceylon; total length, 1,130 mm.

Taylor: The Snakes of Ceylon

0/ i

u

3fi— pn

578 The University Science Bl'lletin

from the ground. The second was obtained at night crawling along
the foundation of a building.

The specimens seem to be placed properly in this species. The
young specimen, No. 30674, was pinkish-fawn in life with diagonal
grayish markings on the sides. On the median dorsal surface are
small deep black spots tending to become paired, although occa-
sionally they are distinctly alternating. A dim dark line is present
from the eye to the mouth angle. The belly is light, peppered
sparsely with grayish, occasionally tending to form spots on the
outer ventrals, in the latter hall' of body.

Boiga ceylonensis (Giinther)

Dipsadomorphus ceulonensis Giintlier, Catalogue of the Colubrine Snakes in the Collection of
the British Museum, 1858, p. 170 (type locality, Ceylon).

Boiga ceylonensis Smith. Fauna of British India, Ceylon and Burma . . .; Reptilia and
Amphibia, vol. 3, Serpentes, 1943, pp. 3.51-353.

This species occurs in India as well as in Ceylon. No specimens
are in the collections.

Boiga forsteni (Dumeril, Bibron and Dumeril)

PI. XXII. lig. 2

Triglyphodon forsteni Dumeril, Bil>ron and Dumeril. Erpetolojiie Generale, vol. 7, 18.54
pp. 1077-1078 (type locality unknown).

Dipsas forsteni Gunther, Reptiles of British India, 1804, p. 30!) ; Anderson, Proc. Zo;,i. Soc
London, 1871, p. 187; Boulenger Fauna British India . . ., 1890, p. 302.

Boiga forsteni Wall, Ophidia Taprobanica or the Snakes of Ceylon. 1921, p. 285; Journ.
Bombay Nat. Hist. Soc, 29, 1924, p. 874; Smith, Fauna (if British India . . .; Reptilia
and Amphibia, vol. 3, Serpentes, 1943, pp. 358-359.

Dipsas forsteni var. ceylonensis Anderson, Proc. Zool. Soc. London. 1871, p. 187 (type
locality, Ceylon).

Two specimens (EHT-HMS Nos. 30676, 30677) of this species
were collected 12 miles north of Trincomalee, Ceylon. Both were
obtained in the immediate vicinity of dwellings at night.

The color pattern of the larger specimen is complex. There is a
series of dorsal light spots, and a second series alternating with these
on the median lateral region. A third series is present still lower on
the side which alternates with the second series. On the edges of
the ventrals is a fourth series. Anteriorly, the spots of the four
series are larger and often contiguous; posteriorly they are nar-
rower and usually separated. Occupying the intervening spaces
are areas or spots of broAvn, many scales of which are black-edged.
These spots arc usually contiguous. On the ventrals are series of

Pl.we XXIIl. Fin. 1- Dri/ophis jmhwndentuti Duincril, Bibron and
Dumeril. EHT-HMS No. 31245 <? ; 12 niilos north of Trincomalee,
Eastern Province. Ceylon: total lonsth, 1,322 mm. Fi^. 2. Microcep-
hnloplvH gracilis Shaw. EHT-HMS No. 30683 5; 12 miles north of
Trincomalee, Eastern Province, Ceylon; total length, 760 mm.

Taylor: The Snakes of Ceylon

579

l^-<

580 The University Science Bulletin

small dark spots. Two larger spots on the outer edges of the adjoin-
ing scales alternate with two or three spots on scales that are con-
tiguous. The small spots are farther from the outer edges of the
scales, and less distinct. A narrow light-edged stripe runs from the
frontal to the nape. A broad brown stripe with a black edge runs
from the eye to the angle of the jaw. All labials are edged with dark
brown.

The scale counts for No. 30676 and No. 30677 are respectively:
Ventrals 264, 257; subcaudals in, 110; scale formula 39-27-29-17.
37-27-28-17; upper labials 9, 10.

Genus Dryophis Dalman

Dri/iiphlx D:iliiuin, Annlect. Entoniol., 1823, p. 7 (substitute name for Dnjimi.'' Menem
[preoccupied] ).

Ge!iot\]ie, Coluber mixutuii Merreni.

Two forms, a brilliant green species and a dull brownish to grayish
species, occur in Ceylon. They are probably equally numerous;
however, the green form is more frequently taken probably due to its
more conspicuous coloration.

Key to the Ceylonese Species of Dryophis

1. Dermal appendage formed of rostral scale only. Green in color rmsutiis

Ventral surface of the dermal appendage formed by rostral, the upper surface
covered with small scales ; color brownish or gray with some transverse black
marks pulverulentus

Dryophis pulverulentus (Dumeril, Bibron and Dumeril)

PI. XXIII, fig. 1

Dryiiuis pulveruleiitu.i Dumeril, Bibron and Dumeril, Erpetologie Generale, vol. 7, pt. 1,
18.54, p. 812 (type locality not known).

Dryophis pulverulcritii'i Boulenger, The Fauna of British India . . ., 189(1, p. 871:
Smith, The Fauna of British India . . ., Reptilia and Amphibia, vol. 3, Serpentes, 1!»43,
p. 378.

Pasxerita purpuraKcens Giinther, Reptiles of British India, p. 307, pi. 23, F (Ceylon).

This species is represented in the collection by five specimens,
EHT-HMS Nos. 30731-30735, all from twelve miles north of Trin-
comalee, Ceylon.

Smith, he. cit., gives a lower average of ventrals and subcaudals
for Ceylon specimens than for Indian specimens. His counts are,
ventrals 179-193; subcaudals 151-178, Ceylon; ventrals 182-203;
subcaudals 169-208 for south India.

The smooth scales are arranged in ol^Hque transverse lines, tiie
scale formula being 15-15-13, the median row being enlarged
slightly. The dermal appendage on the tip of the snout is as long
as the eye diameter.

The general color is brownish when the epidermis is intact or

Taylor: The Snakes of Ceylon 581

grayish with epidermal scales removed. A series of more or less
distinct darker crossbands are present, becoming indistinct poster-
iorly on body. These spots are formed by the darker edges of cer-
tain scales and the black intervening skin. There is a dark cephalic
spot from the prefrontals to the back end of the parietals. An ir-
regularly-edged dark stripe from the anterior end of the first labial
to a point just behind the jaw angle. The ventral surfaces are
powdered with brown or gray.

The following data gives certain scale variations for the species:

Table of Data on Dryophis padveruloitns:

Sub-

Pie-

Post-

Supra -

Infra -

Xiinibor

Sex

Ventials

caudals

oetilar

oculais

labials

lal)ials

30731

S

183

173

3-3

2-2

8-8

9-9

30732

9

181

161

3-3

2-2

8-8

10-10

30733

6

178

165

3-3

2-2

8-8

9-9

30734

c?

183

179

3-3

2-2

8-8

9-9

30735

6

188

175

3-3

2-2

8-8

9-9

Dryophis nasutus (Lacepede)

Coluber nasutus Lacepede, Hstoire Naturelle des Serpents, \ul. 1, p. 100, vol. i. p. 'iT?.
111. 4. fig. ? (type localit.v, Ce.vlon, Guinea, C'arohna. Here restricted to Ce.vlon).

JJryophis myrtemaiis rhodoiiotus Wall, Ophidia Taprobanica or the Snakes of Ce.vlon, 1021,
p. 293 ("a beautiful rose-pink dorsalh' with pinkish-buff \-entrally.'' Galatura Estate, Cejdon,
.OOO-LOOO ft. high).

Dri/opliis nasutus Snulh. The Fauna of British India . . ., Reptilia and Amphibia.
V(j1. 3. Serpentes, 1943, pp. 370-378.

The specimens in the collection are U. S. N. M. Nos. 29428 -Cey-
lon", and 56419-56420 "Ceylon"; EHT-HIMS Nos. 30714-30723, 12
to 15 miles north of Trincomalee, Ceylon. This series has represen-
tatives of several color varieties: yellow-green, grayish green, dark
green, and lavender. All have the two cream lines on the ventrals
and subcaudals, while only a few have also a median pair of white
lines. In three cases these white lines continue to the vent ; in the
others they may be short or they may be absent, or at least not dis-
cernible.

Genus Hurria Daudin

Hurria Daudin, Bull. See. Phihm. Paris, vol. 3, no. 72, Mar.. 1803. p. 187.
Cli'uotype, Coluber cerberus.

It seems that this name takes precedence over Cerberus Cuvier
(1829 Reg. Anim.), which is used for this species by many herpe-
tologists including Smith, loc. cit. See Stejneger, Herpetology of
Japan, Bull. 58, 1907, U. S. N. M., p. 304, regarding the status of
the two names.

A single species, Hurria rhynchops (Schneider), is recognized as
occurring in Ceylon. It is widespread in southern and eastern Asia,
Malay Archipelago and the Philipi)ines.

582 The University Science Bulletin

Hurria rhi/7}cJwps (Schneider I

HiidriiK ri/nrlinpn Sfhneider, Histoiia Amphiliini uin iiaturalis ct literaria, vol. 1, 1799,
p. 246.

Cerhrriis rhi/iirlinps Giintlior. Reptile.^; of Britisli India. isr>4, p. 279.

A specimen (EHT-HMS No. 30682) was captured in a salt water
lagoon 13 miles north of Trincomalee, Ceylon. The color is lead
with very indistinct darker banding; the ventrals are 147; anal
divided; subcaudals 64. The scale formula is (40-27) -23-17. The
keels are very strong on the scales, extending to the outermost row.
(The outer row is not keeled in a Sumatran specimen at hand.) The
labials are atypical in that only the first five are complete, the next
five being divided transversely, leaving a series of very large "tem-
porals" between the labials and the normal temporals. The frontal,
like the parietals, is broken into a number of scales. The greater
part of the three outer scale rows is somewhat pinkish white, form-
ing a stripe, not visible from dorsal view, since three rows are largely
on the ventral surface. The specimen has a single preocular (2 in
the Sumatran specimen).

The species is a common one in river mouths and lagoons es-
pecially near mangrove associations. It also is reputed to enter
fresh water.

Family ELAPIDAE

There are three genera of the Elapidae represented in the island
of Ceylon. They may be recognized by the following key appli-
cable to Ceylon specimens. All are poisonous and among the more
dangerous snakes.

Ki-:y to thk Ceylonese Genkka of the El.apidae

1. Maxillary Ixiiie not extending forward beyond tlie palatine; scales not oblique.

the vertebral .series strongly enlarged ; neck lacking I'longate ribs, and "liood,"

Biaififiriis
Maxillar\- bune extinding forward beyond tlie jialatine; \ertebral series not en-
larged 2

2. Scales in 13-15 rows throughout the body; scales not oblicpie; neck not dilatable,

Callophis
Scales in 15-25 rows on body disposed obliquely; neck with elongate libs, dilat-
able, forming a "liood" -^ "in

Genus Bungarus Daudin

Biiiiyarua Daudin, Mag. PZncycl., vol. 5, Year S (= 1803), p. 434.
Genotype, fa>:ci(itiis.

Two species occur in Ceylon.

Key to Species of Bixgaki's in Ceylon

1. T^iistrous l)lack with a series of paired linear white marks crossing the bod\- :

lower parts white; \entrals, 195-225; subcaudals, 37-50 r(U'riilciis

Black above with Iroui 15 to 21 white crossbars, which are narrow on the vc-tebral
line and widen laterally; may be indistinct in adult; belly white in young;
banded black and white in adult ; ventrals, 21!»-230 ; subcaudals, 32-42, reylonicus*

* This species was inadvertently omitted from the text. No specimens are at hand.

Taylor: The Snakes of Ceylon 583

Bungarus caeruleus (Schneider)

I'ticiidohoa caerulea Schneider, Historia Amphihioruin naturalis et liteiaria, 1799-1801, p.
284 (Based on Russell's An Account of Indian Serpents . . ., vol. 1, p. 2, pi. 1) (type
locality, Vizagapatam, India).

Bungarus caeruleus (part) Boulenger, Fauna of British India . . ., Reptilia and Batrachia,
1890, p. 388; Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921, pp. 437-451.

One specimen is at hand, KUMNH No. 24140, taken at Negombo,
Western Province, Ceylon, by W. C. Osman-Hill.

The specimen is young, measuring 321 mm. in total length; the
tail 36 mm.

The head is dark above, light on supralabials; a partial white
band across back of head. The general color of body is gray-brown
crossed by about 41 narrow transverse cream bands widening a little
low on sides. At the anterior end of the body these are rather
widely separated; at the fourth there is a pair of light lines sep-
arated by one or two scale rows that cross the body separating
wider intervening dark blotches; farther back these white lines are
farther apart until the white bands are separated by distances ap-
proximately equal to the intervening dark blotches. However they
differ from the intervening bands in having the vertebral scales, or
at least some part of the vertebrals, cream in color. The venter is
uniformly white. The bands are obsolete on the tail. The ventrals
are 208; the subcaudals 41, all single save one or two pairs near the
extreme tip. The angle is single. The scale formula is 19 (17) -15-15.

The head has been partially destroyed, presumably by ants, and
all the characters cannot be determined.

Genus Callophis Gray

Callopliifi Gray, Illustrations ot Indian Znolofiy. vul. 2, 1834, pi. 80, fig. 1 (type CaUopliis
gracilii<).

Cullopliis Giinther, Pnic. Znul. Soc. London, 18.')ii, )). 7!».

Only a single species is known from Ceylon,
s Callophis melanin' us (Shaw)

Coluber melaiiurus Shaw, General Zoologj-, vol. 3, 1802, p. 552 (based on Russell's Indian
Serpents, vol. 1, 1790, p. 12, pi. 8) (type locality, Nerva, Bengal); Smith, Fauna of British
India, Ceylon and Burma . . ., Reptilia ard Amphibia, vol. 8, Serpentes, 1943, p. 420.

Callophis trimaculata Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921, pp. 497-
501, fig. 90.

Two specimens of this rare snake were obtained by H. G. Diegnan
at Clodagh Estate, Rattota, Matale District, Ceylon (U. S. National
Museum Nos. 120334, 1203:35. Wall reports 7 other specimens
from the island.

Description: Rostral one third or more, wider than long, the part
visible above about two thirds of the prefrontal suture; internasals

584 The University Science Bulletin

one third wider than long, their mutual suture less than half the pre-
frontal suture; prefrontals slightly longer than wide; frontal length
four fifths of the distance from tip of snout; parietals very large,
their length greater than their distance from the end of the snout,
and twice as great as their width; anterior seminasal higher than
wide, almost inclosing the nostril, much larger than posterior; pre-
ocular very large, longer than high; supraocular rather small, one
half or less the area of the anterior temporal, which touches only
the lower of two postoculars; six supralabials, the order of size
being 1, 2, 4, 5, 3, 6, three scales touching the nasals, third and
fourth entering orbit, fifth touching lower postocular, sixth as long
as the temporal, but narrower; an enlarged secondary temporal also
borders the parietal.

Diameter of eye a little less than its distance from the com-
misure of mouth. Mental very little narrower than rostral; anterior
chinshields one fourth longer than posterior chinshields ; six infrala-
bials, first four touching the anterior chinshield, the fourth very
large; posterior cliinshields separated from the first ventral by four
scales.

Thirteen scale rows from head to anus; seven or eight about base
of tail. Ventrals 231, the first divided; 28 divided subcaudals; anal
divided; no apical pits; under magnification scales appear finely
striated. Maxillary wath a single fang, but no other teeth apparent.

Coloration. Above generally gray-brown with 19 transverse
bands of black on the dorsum, these sometimes complete, sometimes
broken mesially and the opposite half occasionally displaced. The
spot on neck somewhat heavier than others; two black bands en-
circle the tail; the ventral surface bluish gray with six small black
spots arranged more or less symmetrically; venter, fawn on under
side of neck, gradually becoming darker salmon orange to pinkish
on later fifth of body; outer one or two scale rows somewhat fawn,
merging into dorsal and ventral color.

Top of head nearly uniform brownish. A diffuse blackish spot
about eye, extending along lower part of labials to a blackish spot
near angle of mouth but enclosing a roundish cream spot which
covers parts of three scales, but is chiefly on the anterior temporal;
this followed by a lateral cream area extending from throat; nuchal
dark band curving back is more or less connected with dark area at
angle of mouth. Occasional dark flecks, usually confined to median
row or third outer row.

A young specimen, U. S. N. M. No. 120334, from the same local-

Taylor: The Snakes of Ceylon 585

ity agrees in almost all details of squamation. The posterior chin-
shields arc proportionally larger, nearly equalling the size of the
anterior. There are 223 ventrals, and 37 subcaudals. There are 16
dorsal transverse body spots (some broken) with two on tail, both
of which, are interrupted ventrally. The colors are faded. The spot-
ting on the third outer scale row is a little more regular and there
are more of the scattered dark flecks. The small subcaudal spots
are not evident. The flecks on the dorsal scales tend to form lines.
The head is a very light brown with only minute flecks. The eye
spot and the spot at the angle of the mouth are very diffuse, scarcely
discernible without a lens. The neck band is black.

There are certain obvious differences between the specimens here
described and descriptions given of Indian specimens. Thus there
are marked differences in head coloration, the type and others are
described as having a black head and neck; with a pair of yellow
spots on the occiput. The Ceylon adult loses the series of lines on
the back and they are scarcely evident in the young; Smith states
that the under part of the tail is red, while Wall, perhaps speaking
of Ceylon forms mentions the color as pearly gray or pale bluish.
The Indian species is regarded as small (about a foot long). The
larger of these two specimens is approximately 437 mm. (api)rox-
imately llYz inches) suggesting that it is a larger form than the
Indian species. Moreover there is a smaller number of ventrals
(when sexes are compared) in the island specimens. While I believe
the Ceylonese form worthy of subspccific designation, material as
yet is too scanty to delimit the form.

Genus Naja Laurenti

Xnjn Laurenti, .Specinitii UH'dicuni exhiljens Synopsin Reptiliiiin eiiieiidatiiiii, 17CS, p. 00.
Genotype: Coluber vnjn Linnaeus.

Naja naja naja (Linnaeus)

PI. XXIV, fig. 2

Coluber naja Linnaeus, Systema Naturae, 10th ecL, 1758, p. 221. (Based cm Seha, Tlies-
aurus, vol. 1. 1734, pis. 44, 85, 89, 90, 97 [part.]) (type locality, India).

Xaja naja polyocellata Deraniyagala, Ceylon .lourn. Sci., sec. B, vol. 21, l!»;i9, )i. 238 (t\'pe
locality, Polonnaruva, N. Central Province, Ceylon).

Xaja naja naja Smith, The Fauna of British India . . ., Reptilia and .\niiiliiliia. \ol. 3,
Serpentes, 1943, pp. 428, 431; Deraniyagala, Spolia Zeyhmica, \ol. 24, iit. 2, 104.'>. \>. 109.

Deraniyagala Hoc. cit.) has limited the distribution of the sub-
species Naja naja naja to the island of Ceylon, and has applied
names to certain Indian forms. While it is almost certain that there
are forms worthy of names occurring on the continent, Deraniyagala
suggests that since he has not been able to consult the "various rare

586 The University Science Bulletin

old publications containing the different descriptions of Naja" some
of the names may not be correctly applied by him. In view of the
possibility that the name hitescens, which he applied to a penin-
sular race, may be applicable to a northern race, he suggests an al-
ternative name, Naja naja madrasiensis for the same. Smith, loc.
cit., regards Naja naja naja as extending throughout much of penin-
sular India.

The ciiief structural character offered by Deraniyagala in sep-
aration of the Ceylon forms seems to be in the position of the open-
ing of the poison groove. In N. n. naja it is well above the apex,
in the nearest continental form the outlet is said to open at the apex.
The markings differ in details.

There are four specimens in the collections from Ceylon: U.S.N.M.
No. 120333, Peradeniya, Kandy District, Ceylon; EHT-HMS Nos.
30702-30704, 12 miles north of Trincomalee, Ceylon.

All are the tyjiical "spectacled" cobras. They vary somewhat in
the number and distinctness of the black bars across the venter,
which may or may not be traceable across the dorsum and the mark-
ings vary with age. The scale characters of the four specimens are
recorded in the following table.

Table of Data on Ceylonese Specimens of Naja naja naja

Siib-

Supra -

Infra -

Xiilnlii'i'

Sex

Ventials

caudals

labials

labials

Scale rows

30702

6

185

57

7-7

9-9

35. 23, 23, 15

30703

9

187

59

7-7

8-8

35. 23. 23. 15

30704

9

192

60

7-7

8-8

35, 23, 23, 15

120333

9

188

60

7-7

8-8

33, 23, 23, 15

Three first labials touch the posterior nasal; the temporals are 2-3
(in one specimen a small scale makes a count of 2-4). In No. 30702
there is a very small triangular labial between the fourth and fifth
labials. It presumably is a segmented portion from the fourth.

The number of black marks across the venter can be determined
with difficulty in older specimens. They vary in width ; the first and
third or first, second and third being usually widest.

FAMILY HYDROPHIIDAE

Ceylon has known representatives of several genera of the sea
snakes, and it is ])robable that others, not yet reported also occur.
Wall, in his Ophidia Taprobanica, has listed 22 species under 16

Plate XXIV. Fi<i. 1. Agkistrodon hypnale (Merrcm). EHT-HMS No.
30691 9 ; 12 miles north of Trincomalee. Eastern Province, Cevlon ; total length,
354 mm. Fig. 2. Naja naja naja (Linnaeus). EHT-HMS No. 30702 5 ; 12
miles north of Trincomalee, Eastern Province. Cevlon; total length, 936 mm.

Taylor: The Snakes of Ceylon

587

:si^ ^ «;'

588 The University Science Bulletin

ei

genera. 1 have iound authentic records of only the following genera:
Kerilia, Hydrophis, Lapemis, Astrotia, PelamiK and Microcepha-
lophis.

These may be distinguished by the following key.

Key to Ceylonese Genera of' the Family Hyurophudae

1. ^'entials di.stinct throughout length, nonnally entire, not di\ideil 2

Ventrals, except near anterior part of venter, either divided medially by a longi-
tudinal fissure or vestigial (smaller than adjacent scales or absent) 3

2. Scales normal, imbricating, not more than 2.3 rows around the body, nul iiuad-

rangular or juxtaposed Kerilia

Scales tending to be quadrangular, or hexagonal, juxtaposed, not or scarcely im-
bricating ; more than 25 scales around body Hydroplih

3. Head very small, the greatest diameter of body four or five times that of the

neck; the neck and anterior third or two fifths of body very slender, posterior

part compressed and deepened Microcephalopbis

Head not especially small, the diameter of the neck one third or less that of the

greatest body diameter 4

4. Dorsal scales pointed strongly imbricate: ventrals divided into two halves; large

species, of large girth Astrotia

Dorsal scales juxtaposed, subquadrangular in shape 5

5. Ventrals when distinct with a longitudinal fissure; dorsal coloration nearly uni-

form; lateral and ventral coloration lighter (whitish yellow) with or without a
longitudinal darker or lighter lateral line; lower scale rows not larger than others,

Pelamis
Ventrals when present more or less divided ; lower three or four rows of lateral

body scales larger than others Lapemis

Genus Kerilia Gray

Kerilia Gray, Catalogue of the Snakes in the British Museum, 1849, p. !'>7.
Gcnotyi^e, Kerilia jerdoiii.

Only a single species known.

Kerilia jerdoni jerdoni Gray

Kerilia jerdoni Gr?y, Catalogue of the Snakes in the British Museum, 1S49, p. 57 (type
locality, Madras); Smith, The Fauna of British India . . ., Reptilia and Anjphibia, vol.
3, Serpentes, 1943, pp. 440-447, fig. 143.

There is no specimen present in the collection.

This species of sea snake may be diagnosed by the following
characters: large imbricating scales; head shields of the normal
number except loreal absent; scales in 19-23 rows; one preocular,
one postocular; ventrals narrow; six supralabials, anterior temporal,
one; ventrals 225-253. Body of equal diameter throughout. Dark
bantls encircle the body in the young and may be retained in adults.

Genus Hydrophis Latreille

Ilydrophis Latreille, Histoiio Naturelie dcs Reptiles, vol. 4, 1802.
Genotype, faariatui-.

Only three of the numerous species of this genus have been re-
ported in Ceylon. These are cyanocinctus, bitvbcrculatus and
lapemoides.

Taylor: The Snakes of Ceylon 589

These three forms may be distinguished by tlie following synopsis.
Key to Cbylonbse Species of Hydrophis

1. Scales on (he deejiest part of the body with rounded or indistinctl\- pointed tips,

distinctly or feebly imbricate; 5-8 solid maxillary teeth behind the fangs;

body elongate -

Scales on thickest pait of bcdy subquadrangular or hexagonal in shape, juxta-
posed, noninibricate; 8 to 18 maxillary teeth behind fangs; two anterior tem-
porals; 29-3.) scale rows on the neck; ventrals, 314-372; head with a curved
yellow Uiark in young, lost or indistinct in adults lapemoides

2. One anterior temporal; body uniform gray 7 or 8 solid maxillary teeth behind

fangs; greatest diameter of head four times that of neck; scales, 27-48;

ventrals, 276 bituberculatiix

Two anterior temporals; body banded with annuli ; 5 or 6 solid maxillary teeth
behind fangs; diameter of head at most two and one-half times that of neck,

cyaiiochictus .

Hydrophis cyanocinctiis Daudin

Hydrophis cyanorinctus Daudin, Histoire Naturelle des Reptiles, vol. 7, 1803, p. 383 (Based
on Russe'l's .\n Account of Indian SeriJent.", vol. 2, p. 10. pi. 9) (Sandarbans).

See characters in Key. No specimens in tlie collection.
Hydrophis bituberculatus Peters

Hydrophis hituberriilat iis Peters, f>Ionatsl>. Akad. Wiss. Berlin. 1872, p. STt5 (type locality,
Colombo, Ceylon).

No specimens in the collection. The species is known from a single
specimen.

Hydrophis lapemoides (Gray)

Atuira lapemoides Gray, Catalogue of the Specimens of Snakes in the Collection of the
British Museum, 1849, p. 46 (type locality, Ceylon, Madras. Here restricted to Ceylon).
Hydrophis lapemoides Smith, Monograph of the .Sea !^nakes, 1926, p. 86.

No specimens in the collection.

Genus Microcephalophls Lesson

Microcephalophis Lesson, In Belanger, ^'oy. Ind. Orient, 1834, p. 320.
Genotype, gracilis.

Of the two species recognized, only one has been recorded from
Ceylon.

Key to the Species of Microceph.\lophis

Prefrontal not touching the third supralaliial : ventrals. 220-3r)0 (jrarilis

Prefrontal tnuciiing the third supralaijial ; ventrals, 404-468 cantoris

Microcephalophis gracilis gracilis (Shaw)

PI. XXIII, fig. 2

Hydras gracilis Shaw, General Zoology, vol. 3, 1802, p. 560.

Microcephalophis gracilis Wall, Ophidia Taprobanica or the Snakes of Cexlon, 1921. jip.
325-330, fig. 62; Smith, Monogaph of the Sea-snakes (Hydrophiidae), Nov. 27. 1926. pp.
121-213, fig. 34, skull; and Fauna British India including Ceylon and Huiiiin. etc.: Ht'|itilcs
and .\mphibia, vol. 3. Serpentes, pp. 32.')-330, fig. 62, A-V.

I collected this species from the beach 12 miles north of Trincoma-
lee, Ceylon (EHT-HMS No. 30083). The specimen was very active

590 The University Science Bulletin

but wholly unable to make any progression on the wet sand. It had
been stranded by a small wave, and when found was less than four
feet from the edge of the water.

This species is regarded as rare thruugliuut must of the wide area
where it occurs. Smith (Monograph) reports a Ceylon specimen,
and Wall {loc. cit.) mentions four specimens in the Colombo Mu-
seum, presumably from Ceylon. He reports it as common on the
Malabar and Coromandel Coasts of India, and gives its range as
extending from the Persian Gulf to Japan. An Australian record
needs further confirmation since it has not been recorded in the in-
tervening territory. The other species of the genus (iV. cantoris)
is unknown as yet in Ceylonese waters.

The arrangement of head scales conforms to Wall's figure save that
the rostral is higher and the width of the rostral one third greater. The
ventral count is 233, the subcaudals 38. The latter part of the body is
extremely compressed, the dorsal part forming a sharp crest. The
anterior narrowed part however is nearly cylindrical. The depth of
the head at the mouth angle (5.3 mm.) is contained in the greatest
body depth (22 nun.) about four times. There are 31 scales around
the neck (33 counting the divided ventrals), and 29 a centimeter in
front of the anus.

Several small barnacles are fastened to the scales of the specimen.

Smith has recognized a subspecies of gracilis on the coast of the
Malayan Peninsula and Java.

Genus Astrotia Fischer

Astrotia Fischer, Abli. Nnturw. Vei-. Uaiiiljinc. \(il. 8, ISoli. p. 38.
Genotype, si-hizopholis = stokefi.

Only a single species known.

Astrotia stokcsii (Gray)

Hydrus stokesii Gray, In Stokes' Discovery of Australia, vol. 1, p. 502, pi. 3.

Astrotia stokesii Wall, Mem. Asiat. Soc. Bengal, vol. 2, 1909, p. 250; Ophidia Tapro-
banica . . ., 1921, p. 396; Smith, The Fauna of British India . . ., Reptilia and
Amphibia, vol. 3, Serpentes, 1943, pp. 471-472.

This, the largest of the sea ser])cnts, is known from the western
coast of Ceylon. No specimens are in the collection.

This form may be recognized by the character of the ventral
scales. These are divided in two, except anteriorly, and the halves
are pointed or with a dentate tip. The ventrals are 226-286, the
preanals strongly enlarged. The diameter of the neck is more than
half the greatest body diameter. One preocular and two postoculars
are present. The supralabials are 8-10, the infralabials 10-12. The

Taylor: The Snakes of Ceylon 591

scale ro\^•s ^•al•y between 37 and 47 on the neck, and 47-59 on the
body.

The color is "yellowish or pale brown, with broad black or dark
brown bands more or less complete, or with dorsal bars and ventral
spots. Spots or narrow bars often present between the annuli.
Head dark olivaceus to yellowish."

The species reaches a length of at least 1600 mm. and the girth
of a large specimen may exceed 260 mm.

Genus Pelamis Daudin

Pehnius (in part) Diiiidln, Histoire Naturelle, Generale et Particuliere des Reptiles (1S03),
Year XI, vol. VII, pp. 357, 361, 3(32.
Genotype, Aiiguis platurux.

Only one species is generally recognized. It is one of the most
widely distributed of all snakes. There are a number of so-called
color varieties.

Pehimis i)lntiiriis (Linne)

Aiiguis platurus Linne, Systema Naturae, ed 12, 1766, p. 391.

Pelamis platurus Gray, Ann. Philos., 1825, p. 15; Smith, Monograph of the Sea-snakes
(Hydrophiidae), Nov. 27. 1926, pp. 116-120, fig. 33; Smith and Taylor, U. S. Nat. Mus. Bull.,
No. 187. 1945, pp. 176-177.

The puzzling variety of color i)atterns and the remarkable varia-
tion in scale rows and ventral scale counts suggest strongly that the
assemblage of forms now placed under this specific designation rep-
resents more than a single species. The scale rows reputedly vary
from 49-67 rows, a difference of 18 rows. The ventral scales vary
between 264-406, a difference of 142. Few if any other snakes have
scale variations of this magnitude.

Specimens are at hand from Ceylon belonging to two or three
well-marked color forms.

U. S. Nat. Mus. No. 19213 is a specimen in which the body is
blackish above (faded), and yellowish below, the black beginning
on the head in the region of the nostrils and extending along the dor-
sum in a band 11 scale rows in width, for about one third the length
of the body. Where this ends, there begins a series of transverse
bands, contiguous mesially and extending down on the sides. Pos-
teriorly the dorsal parts are separated. There are no lateral sj^ots
save on the tail where small spots intervene on the sides between the
bands.

U. S. Nat. Mus. No. 31212 has the black coloration forming a
series of spots contiguous on the median dorsal line. Posteriorly the
spots are rhomboidal, extending rarely half way down the side. The
intercalated spots may be distinguished by their being lower on the

592 The University Science Bulletin

sides of the tail. This specimen, although a shorter specimen than
the preceding one. has the nostrils distinctly farther back from the
tip of the snout.

A third specimen from the northeast coast, 12 miles north of
Trincomalee, is coal-black on the dorsal 25 scale rows, and dirty,
grayish lavender on the ventral part of the body save for an indis-
tinct lighter line separating the two colors. The labials are not
lighter than the ventral coloration. The tail is clear pinkish white
with a series of transverse coal-black bands, contiguous or not, dor-
sally, and a series of black spots alternating with them on each side.
Tn front of and behind the vent there is a deep black spot.

The ventral-subcaudal scale counts for Nos. 31213, 31212 and
31250 are, respectively, 334-44, 369-52, 360-55; and the midbody
scale rows are, respectively, 58, 51, 58. The four preanals in No.
31212 are reduced to the size of other ventral scales and the scales
preceding these are not enlarged.

Thus if one considers these variant forms as worthy of name<,
they must be regarded as species rather than subspecies.

Genus Lapemis Gray

Lapemis Gia>-. Ilhistiati(jus of Indian Zoulogy, chiefly selected fiuiii the collection of Majui-
Geneial Hardwicke, vol. 2, 1S3.5, pi. S', fig. 2.
Genotype, Lapemis curtus Shaw.

Two species are known.

Lapemis curtus Shaw

Lapemis curtus Siiaw. General Zoology, vol. 3, 1802. p. ."102 (type locality not given).

A single specimen, EHT-HMS No. 30678 $ , was taken on the
shore of the Bay of Bengal, 13 miles north of Trincomalee. The
following scale characters obtain: frontal shorter than its distance
from rostral; parietal broken into three or four unequal scales; one
preocular, one postocular; seven supralabials, the two last very
small; second supralabial bordering the prefrontal; fourth only bor-
dering orbit; two large anterior temporals, two posterior temporals;
three infralabials border the chinshields; both pairs of chinshields
separated by one or two scales.

The scale rows counted at various points on body arc 35-37-30-
40-33; scales more or less quadrangular, juxtaposed; ventrals 195,
more or less distinct throughout; subcaudals 45 (scales in a row
on lower edge of tail).

Head normal, the body greatly thickened and laterally com-
pressed; at its greatest depth approximately three times depth of

Taylor: The Snakes of Ceylon 593

neck; back with a moderately sharp ridge; gray with oS bands
across the back separated by one or two scales on the median dorsal
line and coming to a point low on side; below these and on venter
yellowish buff.

Family VIPERIDAE
The family consists of two subfamilies, Viperinae and Crotalinae.

Subfamily Viperinae

This subfamily is represented in Ceylon by two genera, Echis and
Vipera, each represented by a single species.

Genus Vipera Laurenti

Vipera Laurenti (part). Si)pciiiit'n iiu'diciiiii, exliili<>ns Synniisin Reptiliuiii I'liii'iulatuin. 1708,
p. 99.

Genotype, redi = aspis.

Vipera russelli russelli (Shaw)

Cohihrr russelli Shaw, Naturalists Miscellany vol. 8, pi. 291 (1707) (t\-pe localit>', India,
l)y inference).

Vipera russelli Strauch, Mem. .\cad. iSt. Petershuris, 7, XH', No. (1, p. 85: Wall. Ophidia
Taprobanica or the Snakes of Cejlon, 1921, p. itOi.

Vipera russelli russelli .Sniilh. The Fauna of British India . . .. Replilia and Amphibia,
vol. S, Serpentes. 1943, p. 483.

Two specimens (U. S. Nat, Mus. No. 115611 and EHT-HMS No.
30679) are at hand, the latter taken 12 miles north of Trincomalee,
Ceylon. I have the skin of a fine specimen (EHT-HjMS No. 31254)
presented to me by Major (Jeneral Richardson. It was killed by
him in his yard in Trincomalee.

This snake is not rare in this part of Ceylon. Several dead speci-
mens were seen along the road.

The species is not aggressive. Usually, when disturbed, it hisses
rather loudly.

Genus Echis Merrem

Echis Merrem (ixirt.). Tcntamen .'^xstematis .\mijhdiinrum, l.s2n. ji. 4!).
Genotype, Eiliis ciiriiiala.

Ecliis carinatiis Schneider

I'seudoboa rariuata Schneider, Historia Aniphibiorum naturalis et literaria, \ol. 2, 1801, p.
285 (species based on Russel, An Account of Indiana .Serpents . . . )

Echis rarinata Fayrer, The Tiianatophidia of India . . ., 1874, pi. 12; Wall, Ophidia
Taprobanica or the Snakes of Ceylon, 1921, pp. ."i29-547, figs. 93-95.

No Ceylonese specimens of this species are in the collection. I
have taken it in the city of Bombay.

In Ceylon the species is known only from the extreme northern
part and may be of relatively recent introduction. It is a small

37—90

594 The University Science Bulletin

snake that has the habit of producing a curious rasping sound. The
scales have a sawlike crest or keel which when rubbed together pro-
duce the sound.

Sl'BFAMily Crotalinae

This subfamily is represented by two genera, Agkistrodon and
Trimeresurus the first with two, the second with one species.

genus trimeresurus Lacepede

Trimeresurus Lacepede, Ann. du Mus. Paris, vol. 4, 1804, pp. 196, 209 (part).
Genotype, Triiiuresuruii virid s.

This genus name as applied to Asiatic forms, is. I believe, com-
posite, as is the name Bothrops as applied to related groups of pit
vipers in Mexico, Central and South America.

Only a single species is known from Ceylon.

Trimeresurus trigonocephalus (Sonnini and Latreille)

Vipera trigonocephala, Sonnini and Latreiile, Histoire Naturelle de.s Reptiles, vol. 3, 1801
(1803?), p. 175 (type based on Seba, Thesaurus, vol. 2, pi. 36, No. 2 [said to be from the
Island of St. Eustace]); Daudin, Histoire Naturelle Generale et Particuliere des Reptiles, vol.
6, year XI (1803), pp. 175-177.

Cophias trigonocephalui iVlerreni, Tentamen Systematis Amphibiorum, 1820, p. 156.

Megaera trigonocephala Wagler, Natiirliches System der Amphibien, 1830, p. 174 ; Gray,
Zoological Miscellany, 1842, p. 49; Kelaart Prodromus Faunae Zeylanicae, being Contributions
to the Zoology of Ceylon, vol. 1, 1853, p. 142; and idem. vol. 2, pt. 7, 1854, p. 8 (trigone-
cephala).

Trigonocephalus iiigro-margiiiatus Kuhl, Beitrage Zool., 1832 (1820), p. 90; Schlegel, Essai
sur la Physionomie de.s Serpens, 1837.

Bothrops nigro-murginatus Dunieril, Bibron and Dunieril, Erpetologie Generale, vol. 7, pt. 2,
pp. 1515-1517.

Megaera olivnrea Gra.\', Zoological Miscellany, 1842, p. 12 (type locality not known); and
Catalogue of the specimens of Snakes in the collection of the British Museum, 1849, p. 12.

Trimeresurus trigouoccphalus Giinther, Reptiles of British India, 1864, p. 390; Theobald,
Descriptive Catalogue of the Reptiles of British India, 1876, p. 223; Boulenger, The Fauna of
British India, including Ceylon and Burma; Reptilia and Batrachia, 1890, p. 431; Werner,
Ahh, Vehr, Zool. Bot. Ges. Wien, vol. 43, 1893, p. 352; and Sitzb. Vehr. Zool. Bot. Ges.
Wien, vol. 46, 1896, p. 10; Abercromby, The Snakes of Ceylon, 1910, pp. 49, 69; and Spoilia
Zeylanica, vol. 7, 1911, p. 207; idem ibid., p. 304; Wall, Ophidia Taprobanica or the Snakes
of Ceylon, 1921, pp. 560-564, fig. 98 (head); Journ. Bombay Nat. Hist. Soc, vol. 30, 1925,
p. 249; and Poisonous Snakes of India, 1928, p. 50; Smith, The Fauna of British India in-
cluding the whole of the Indo-Chinese subregion ; Reptilia and Amphibia, vol. 3, Serpentes
Dec, 1943, pp. 506-507. "'"

Lachesis trigonocephalus Boulenger, Catalogue of the Snakes in the British Museum (Natural
History), vol. 3, 1896, p. 559; Pearle-ss, Spoilia Zeylanica, 1909, p. 54 (Badula).

One specimen (No. 5894) is present in the U. S. National Museum
Collection.

Scale rows are 26-21-19-15; supralabials 10-11; infralabials 12;
one or two pairs of suboculars separate the labials from the eye;
ventrals 147, the terminal ventral divided forming two scales deeply
notched between; anal single; subcaudals, 58, double. The supra-
ocular is divided, and separated from those of the opposite side l)y
five irregular scale rows.

Taylor: The Snakes of Ceylon 595

The skin is dark, approaching coal bhick near the dorsal surface.
Most of the scales are some shade of green with black areas with
elongate extensions. This series of markings may be more or less
connected. Upper part of head greenish, the scales often partially
black-edged. A broad black stripe present from eye to mouth angle.
The supralabials are yellow, with a dark line crossing below eye;
the ventrals are yellowish or greenish, edged with yellowish. The
under side of head is yellowish. The tail is black at the tip.

Genus Agkistrodon Beauvois

Af/klstrodoii Beauvois, Trans, .\inerican Phil. Soo.. vol. 4, 1799, p. 381.
Cienotype, Agkistrodon mokasen.

Two forms of this genus are recognized in Ceylon. These may be
differentiated by the following key.

Key to Ceylon Species of Agkistrodon

A. Hump on snout tip occupies only the middle of the tip; ventrals, 120-135; sub-

caudals, 28-30 ; 380 mm. in total length nepa

B. Hump on snout tip occupies the whole of the tip; ventrals, 138-157; subcaudals,

32-46 ; 545 mm. in total length hypnalc

Agk istrodon n cpa { Laurent i )

Coluber nepa I.iiurenti, Specinien medicum. exhibens Synopsin Reptilium emendatum, 1768,
p. 97 (based on Selia, vol. 1. pi. 19, fig. 7) (type locality, erroneously "Madagascar").

I/ypnale nepa Giinther, Reptiles of British India, 1864, p. 394 (part.).

Anristrodon nepa Smith, Journ. Bombay Nat. Hist. Soc, vol. 39, 1937, p. 730; and Fauna
of British India, Ceylon and Burma including the whole of the Indo-Chinese subregion ;
Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp. 500-501, Ceylon (Hagkalla, Kandy, Ani-
bewela, Mudulkale).

Ancistrodon hypnale Boulenger, Tlie Fauna of British India including Ceylun and Burma,
1>!HI, p. 424 {part.).

Aneistrodon miUardi Wall, Ophidia Taprobanica or the Snakes of Ceylon, 1921, p. 554,
fig. (not miUardi. Wall, 1908); Journ. Bombay Nat. Hist. Soc, vol. 30, 1925, p. 249.

I have examined two specimens, KUMNH Nos. 24142, 24143
(which lack exact locality data). These have the following scale
counts respectively: Ventrals 140-139; subcaudals 42-36; anal
single in both. The scale formula in each is 17-17-15.

The rostral is strongly elevated, at least two fifths extending
above the surface of the snout. The posterior side of the elevation
is covered with some 15 small scales. The area in front of the fron-
tal is occupied by 16 imbricating scales. The frontal, supraoculars,
and parietals are normally large and unbroken. The supralabials
are 7-7; the infralabials 8, 8, with only three touching first chin-
shields. The color above is brown or fawn with indistinct darker
marks on each side that sometimes are fused on the middorsal line.
The venter is thickly peppered with dark pigment.

596 The University Science Bulletin

Agkistrodon hi/puale Mcrroin

PI. XXIV. ««. 1 ; PI. XXV

Cophins liypiialt' Mtireni. Tentamen Systeiiiatis Amphil)ioium, 1820, p. 155.

Triyniiorephalus liypiiale Schlegel, Essai sur le PliysioiKnnie des Serpens, 1837, p. 550. pi. 20,
fig. (type locality, "Ceylon").

Tri))teresurus ? ceylunensif: Gra>', 1S42, Zoological Miscellany, 1842, p. 4!l (Ceylon).

Hypnale nepa Giinther, Reptiles of British India, 1864, p. 394 (part).

Hypiwie nfinia Anderson, .Tonrn. .\siatic Sor. Bengal. 40. pt. 2. 1871, p. 20 (Ceylon).

Aiiostrodon hiipiialc (part) Boulenger, The Fauna of British India including CeyU.n and
l^uinia; Rei>tilia and Batrachia, 1890, pp. 424-425; Catalogue of the Snakes in the British
Museiuii of Natural History, 2d ed., vol. 3, 1896, p. 7G; Wall, Spolia Zeylanica, vol. 3, pt. 10,
Oct.. 1905, pp. 140-147; ibid., vol. 11, 1920, p. 403; ibid., vol. 12, 1924, p. 270; Jovnii.
Bombay Nat. Hist. Soc, vol. 30, 1925, p. 248; Ophidia Taprobanica or the Snakes of Ceylon,
1921, pp. 549-554, fig. 96 (head); Fearless, Spolia Zeylanica, vol. 6, pt. 21, Mar., 1909, p. 55;
.^bercroniliy, ibid., vol. 8, pt. 32, 1913, pp. 304-305 (It is possible some of the references of
Wall, Peerless and Abercromby may refer to A. nepa] ; Smith, The Fauna of British India,
Ceylon and Burma including the whole of the Indo-Chinese subregion ; Reptilia anil Amiih!bia.
vol. 3. Serpentes. 1943. pp. 499-500.

Tlie adult males and females differ considerably in color, that of
the female being very much lighter in shade, with the dark mark-
ings usually more intense and more strongly contrasting.

The general dorsal coloration is lavender-brown with dim or well-
defined brown or blackish markings. The light areas on the back
tend to separate the dark spots. A dark line, more or less distinct
runs from the eye to the angle of the mouth, where it may continue
back along the edge of the ventrals for some distance as a narrow
black line, continuous or more usually broken. A white or cream
line usually borders the upper edge of the dark line on the head and
this may follow along the anterior edge of the ventrals as a series
of broken irregular white fiecks. On the outer scale row alternating
scales usually have small black dots.

Variation in scale counts are recorded in the accompanying lable.

Specimen No. 30753 contained 9 embryos. These were well ad-
vanced and scale counts were made on the lot. The ventrals and
subcaudals are: 146-38, 148-36, 149-36, 150-39, 152-38, 150-36, 151-
38, 149-42, and 149-44. The last two are male specimens and have
the hemipenis extruded. The others are presumably females. Thus
the range for the brood is 146-152 for ventrals; 36-44 for subcaudals.
A single male specimen in the United States National Museum from
an unknown Ceylon locality has 151 ventrals and 44 subcaudals.

In the species the anal scales are invariably undivided, and there
are three preoculars, one subocular and one postocular.

The species was found to be the most common terrestrial snake
in the region about Trincomalee. Specimens were discovered en-
sconsed under logs, under small piles of trash, or crawling about

Taylor: The Snakes of Ceylon

597

-^~ ' -^".^

'<^iS'H^

-^^-

^\

i~

r..

Plate XXV. Agklslidilon hypnalc (Menem). EHT-HMS No. 30687 9;
12 miles north of Tiinroinalee, Ea.'^tein Pio\inco. Ceylon; total length, 434
mm.

598

The University Science Bulletin

during the day or night on the forest floor. They were slow to anger
and only struck when teased or injured. It is not regarded as a
dangerous snake by the natives.

Scale Coi^nts of Agki.^trodon hypnak

HT-HMS

Scale

Upper

Lower

Pre-

No.

Sex

VentritLs

Caudals

foniiula

labials

labials

ocular

30684

9

153

38

20-17-15

7-7

9-9

3-3

30685

9

147

36

19-17-15

7-7

9-9

3-3

30686

$

154

46

19-17-15

7-7

8-8

3-3

30687

i

154

44

19-17-14

7-7

9-8

3-3

30688

A'tt.

153

37

19-17-15

7-7

9-9

3-3

30689

'9

150

39

19-17-15

7-7

9-9

3-3

30690

9

153

38

19-17-15

7-7

9-9

3-3

30691

9

152

38

19-17-15

7-7

9-9

3-3

30692

o

-t-

156

38

19-17-15

3-3

30693

$

149

41

19-17-15

7-7

9-9

3-3

30694

S

153

45

19-17-15

3-3

30695

9

153

38

17-17-15

7-7

9-9

3-3

30750

9

150

37

17-17-15

7-7

9-9

3-3

30751

S

152

45

19-17-15

7-7

8-8

3-3

30752

vs.

153

37

19-17-15

8-8

9-9

3-3

30753

9

146

38

19-17-15

7-7

9-9

3-3

30754

9

153

37

19-17-15

7-7

7-9

3-3

30755

S

154

45

19-17-15

7-7

9-9

3-3

Taylor: The Snakes of Ceylon 599

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1796. An Account of Indian Serpents collected on the ccast of Coromandel,
containing descriptions and drawintrs of each sjiecies, together with
experiments and remarks on their .several poisons. Vol. I, 1796, pp.
1-90, 1-44 colored plates. Vol. II. A continuation of an Account of
Indian Serpents; containing descriptions and figures from .specimens
and drawings transmitted from various parts of India. 1801-1809,
pp. 1-53, 41 colored jilates. Including index of both volumes. I.ondon.

Smith, M.'Vlcolm A.

1926. A monograph (ji the Sea Snakes. 1926, pp. 1-130, text figs. 1-35, pis.
1-2. London.

602 The University Science Bulletin

1943. The Fauna of British India, Ceylon and Burma including the whole
of the Indo-Chinese Sub-region; Reptilia and Amphibia, Vol. 3
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Sp.var, a. E.

1910. The Bile of Ru.^^.sell's Viper. Spolia Zeylanica, vol. 6. pt. 24, Mav.
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S.'iRASIN, F.

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W.ALL, Fr.ANK

1905. Notes on Snakes collected at Hakgalla, Cevlon. Spolia Zeylanica,

vol. 3. pt. 10, Oct., 1905, pp. 144-147.
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vol. 4, pt. 16, Aug., 1907, pp. 166-172.

1907. Occurrence of the Indian Snake Lycodon striatus in Ceylon. Spolia
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1907. The Common Indian Krait, Bungarus candidus, in Cevlon. Spolia
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1907. Remarkable Snake fatalitv. Spolia Zevlanica. vol. 4. pt. 16, Aug..
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1921. Notes on some Ceylon Snakes recentlv acquired bv the Colombo Mu-
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1923. Notes on the distribution of the Snake Aspiduru irachyprocta. Spolia
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Werxer, F.

1893. Bcmerkungen liber Reptilien und Batrachier aus den tropischen Asien

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WiLLEY, A.

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Taylor: The Snakes of Ceylon 603

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1904. Denclmpliis bifrcnnlis Boulenger. Spolia Zevlanica, vol. 1. 1904, i)p.
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1906. Tene.'itrial Colubridae of Cevlon. Spolia Zevlanica, vol. 3, pt. 12,

April, 1906, pp. 227-234.
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1910. As.sociation of barnacles with Snakes and Worms. Spolia Zeylanica,

vol. 6, pt. 24, May, 1910, pp. 180-181, plate.

23-90

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