— fV I \

HARVARD UNIVERSITY

R

LIBRARY

OF THE

Museum of Comparative Zoology

BULLETIN o/-'«

The University of Kansas

SCIENCE BULLETIN

(Continuation of Kansas University Quarterly)

Vol. XXV

LAWRENCE, KANSAS
Published Semimonthly

Vol. 39

JUNE 1, 1938

No. 11

Entered as second-class matter December 29, 1910, at the post office at
Lawrence, Kansas, under act of July 16, 1894

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

Lawrence, Kan.

EDITORIAL BOARD

H. B. HuNGERFORD, Chairman A. W. Davidson

E. H. Taylor, Secretary C. M. Baker

J. D. Stranathan 0. 0. Stoland
R. C. Moore

THE

KANSAS UNIVERSITY

Science Bulletin

DEVOTED TO

o-^

UNIVERSITY OF KANSAS-
Science Bulletin /^^'^'/^.'^

<S9

First Mailing of Volume 25 ]ul^ 10, 1939

LAWRENCE, KANSAS

1938

PRINTED BY KANSAS STATE PRINTING PLANT

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TOPEKA 1938

17-4141

NOTICE TO EXCHANGES

The attention of learned societies and other institutions which
exchange scientific publications' with the University of Kansas is
called to the list of publications of this University on the third and
fourth pages of the cover of this issue.

Those marked "Supply exhausted" cannot be furnished at all ; as
far as the supply permits the remaining numbers will be furnished
gladlv to anv of our exchanges who mav need them to complete their

EDITORIAL BOARD

H. B. HuNGERFORD, Chairman A. W. Davidson

E. H. Taylor, Secretary C. M. Baker

J. D. Stranathan 0. O. Stoland
R. C. Moore

THE

KANSAS UNIVERSITY

Science Bulletin

DEVOTED TO

THE PUBLICATION OF THE RESULTS OF

RESEARCH BY MEMBERS OF THE

UNIVERSITY OF KANSAS

Vol. XXV

(Whole Series, Vol. 35)

PUBLISHED BY THE UNIVERSITY
LAWRENCE, KANSAS

1938

PRINTED BY KANSAS STATE PRINTING PLANT

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S^ f^A - L

IX^SS

CONTENTS OF VOLUME XXV

No. Page

1. A Preliminary Survey of the Fresh-water Algae of Eastern

Kansas. Rufus Thompson 5

2. Effects of Alkaloids on the Growth of Fungi. James C.

Bates 85

3. The Flora of the Sand Hills of Central Kansas. Jacob H.

Doell 113

4. Studies in Anaphylaxis XVI: Physiological Studies of His-

tamine and Peptone Reactions in the Cat. Paul Kabler, 149

5. Studies in Anaphylaxis XVII: Physiological Studies of the

Hypersensitive Cat. Paul Kabler and A^. P. Sherwood. . 159

6. A New Fish, lAstracanthus eliasi, from the Pennsylvanian

of Nodaway County, Missouri. C. W. Hibbard 169

7. Distribution of the Genus Reithrodontomys in Kansas. C.

W. Hibbard 173

8. A Lower Jaw of Martinogale alveodens Hall. D. H.

Dunkle 181

9. Weights and Linear Dimensions of the Skull and of Some of

the Long Bones of the Mourning Dove {Zenaidura ma-
croura carolinensis) . H. B. Latimer and C. W. Asling . . . 187

10. Weights and Linear Dimensions of the Skull and of Some of

the Long Bones of the Red-tailed Hawk {Buteo borealis
borealis) . JI. B. Latimer 199

11. On the Alkyl Derivatives of the Isomeric Ortho and Para-

Phenoxyphenyl Thiazolidones. M. E. Roberts and F. B.
Dains 213

12. Notes on the Snakes of the Genus Salvadora. Hobart M.

Smith 229

13. Miscellaneous Notes on Mexican Snakes. E. H. Taylor and

Hobart M. Smith 239

14. Concerning Mexican Salamanders. E. H. Taylor 259

15. Notes on the Mexican Snakes of the Genus Leptodeira, with

a Proposal of a New Snake Genus, Pseudoleptodeira. E.

H. Taylor 315

(3)

4 The University Science Bulletin

No. Page

16. On Mexican Snakes of the Genera Trimorphodon and Hyp-

siglena. E. H. Taylor 357

17. New Species of Mexican Tailless Amphibia. E. H. Taylor, 385

18. A New Anuran Amphibian from the Pliocene of Kansas.

E. H. Taylor 407

19. Frogs of the Hyla exirnia Group in Mexico, with Descrip-

tions of Two New Species. E. H. Taylor 42]

20. A Contribution to the Taxonomy of the Subfamily Issinae

in America, North of Mexico (Fulgoridae, Homoptera).
Kathleen C. Doering 447

21. The Morphology of the Carolina Mantis. Philip Levereault, 577

22. A Monographic Revision of the North American Species of

Stenelmis (Dryopidae: Coleoptera). Milton W. Sander-
son 635

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 1

A Preliminary Survey of the Fresh-water Algae of

Eastern Kansas

RUFUS H. THOMPSON, Department of Botany, University of Kansas

Abstract: This paper reports 241 species for Kansas. Of this number 202
species are new records for the state and include the descriptions of the fol-
lowing five new forms :

Euqlcnn truncata Walton var. baculifera var. nov.

Euglena alata sp. nov.

Phacus infiata sp. nov.

M erism(ypedia angularis sp. nov.

Cosmarium sexpapillosum sp. nov.

Lepocinclis truncata Da Cunha described from Brazil and Lepocinclis tur-
biniforrnis Deflandre, described from Venezuela, are reported from the United
States for the first time.

INTRODUCTION

PREVIOUS publications on the fresh-water algae of the state of
Kansas have been those of Prof. F. W. Cragin (Bull. Washb.
Coll. Lab. (1:1, 1884), (1:2, 1885), (1:6, 1886), (2:9, 1889), B. B.
Smyth (Trans. Kansas Acad. Sci. 13:1892), G. H. Curtis (ibid. 17:
1899-1900), J. B. McNaught (ibid. 29:1920), L. L. Bailey (ibid. 35:
1932), and S. A. Mannoni (ibid. 35:1932). Collectively, these pub-
lications give a list of 141 species of algae for the state. This list
contains representatives in 5 classes, 5 orders, 8 families and 46
genera.

The algae being reported in the present paper represent 9 classes,
19 orders, 44 families and 116 genera. The four new classes are
Rhodophyceae, Dinophyceae, Cryptophyceae and Charophyta. The
total number of species and varieties is 241. The following is a

(5)

6 The University Science Bulletin

table illustrating the taxonomic distribution of the algae in the
present paper.

Class Orders Families Genera Species

Myxophyceae 3 7 21 39

Rhodophvceae 1 1 1 1

Heterokontae 2 2 2 2

Chrysophyceae 2 3 4 4

Crvptophvceae 1 1 1 1

Clilorophyceae 9 28 79 146

Dinophyceae . . 1 2

Euglenophvceae 1 5 44

Charophyta 1 1 2 2

19 44 116 241

From this table it is easily seen that the Myxophyceae, Chloro-
phyceae and Euglenophyceae comprise the bulk of the determined
species and that the majority are in the Chlorophyceae. Aside from
the fact that the class Chlorophyceae contains the largest number
of species of fresh-water algae of all the classes, its overshadowing
of the other classes in number of species cited in most taxonomic
papers is perhaps also due to three other facts; first, that a large
number of the Chlorophyceae are nonmotile; second, that they
either form macroscopic thalli or aggregate in such amounts as to be
very prominent ; and third, that they themselves are either large
or of a striking morphological distinction.

Although the algae reported in this paper were collected for the
most part within Douglas county, they afford a good representation
of the possible algal flora of the state as a whole. This is true since
they are, with few exceptions, the most common forms met with in
any locality and have in many cases been repeatedly collected from
several different stations. The few collections the author was able
to make in the southern and central portions of the state yielded
some new species. They were for the most part desmids and other
algae of acid waters. This was to be expected, since the collections
were made in sandstone and sand-hill regions. An intensive and
extensive study of the waters in those regions would undoubtedly
yield more new forms and they would probably be largely desmids
and other algae of acid waters.

• In these collections were also found many species that had already
been found in the vicinity of Lawrence. Notable among them, in
the unicellular forms, were species of Trachelomonas, Euglena,
Phacus, Lepocinclis and Tetraedron; in the filamentous forms of
green algae, species of Spirogyra, Zygnema, Cladophora, Pithophora,
Stigeoclonium, Hormidium and Chaetophora; in the unicellular and
filamentous blue-greens, species of Chroococcus, Gleocapsa, Oscil-

Thompson: Fresh-water Algae 7

latoria and Phormidium. These organisms seem to be tolerant to
a change of waters from sUghtly acid to quite alkaline, from an
approximate acidity of pH 6.5 to 7.5.

The algae described in this paper have been collected from in-
termittent pools, swamps, small ponds, lakes, streams, moist soil,
and moist rocks, the majority being from still or slowly nmning
waters. The southern collections were made from the Verdigris
river and a small lake at Neodesha, Chetopa creek at Altoona,
Neosho river, Santa Fe pond. Village creek and an adjacent swamp
at Chanute, and Deer creek at Garnett. Collections were also made
from the temporary waters of ditches and small pools in each of
these places. The collections from central Kansas were made from
a small lake and intermittent pools in the sand hills near Burrton.
At Lawrence the majority of the collections were made from the
boat slough at the north end of Ohio street and the adjacent swamp
by the Kaw river. Other stations from which frequent collections
have been made are at the Kaw river dam, the drainage ditch
south of Haskell Institute, a pond on the property of R. R. Jack-
man about one mile west of the University campus, and an inter-
mittent pool in the abandoned quarry on east Fifteenth street. Col-
lections were also made from the AVakarusa river, the country club
spring, the city park pool, and many temporary waters in ditches
along outlying roads.

There have been quite a number of species of algae in culture
which have not been identified, either because there has not been
sufficient material, or the necessary stages in reproduction have not
been evident for complete determination. For example, there have
been a number of obviously different species of Spirogyra and
Mougeotia in culture, but there have been no fruiting bodies. Since
accurate, specific determination of the members of these two genera
is entirely dependent on the dimensions and characters of the zy-
gotes, or in the case of Mougeotia the aplanospores also, they have
been undetermined. Some of the species of Spirogyra that have
been reported for the state (Mannoni, 1932) have been determined
entirely on the dimensions of the vegetative cells and the number
of chloroplasts within f\ cell. Neither of these is a good criterion
to species without the zygote, since cell measurements vary with
the age of the cell in width as well as in length and the number of
chloroplasts may vary among the cells of the same filament. Mou-
geotia is difficult to determine since fruiting material is very rare
and determination is absolutely dependent on the characters of the
zygote or aplanospore.

8 The University Science Bulletin

Four new species and one new variety are being described in
this paper: Euglena truncata Walton var. bacidijera var. nov.,
Euglena alata sp. nov., Phacus inflata sp. nov., Merismopedia an-
gidaris sp. nov., Cosmarium sexpapillosnm sp. nov. Also, two species
of Lepocinclis are being reported for the United States for the first
time: Lepocinclis truncata Da Cunha and Lepocinclis tiirbiniformis
Deflandre. L. truncata was described and reported from Brazil
only, and L. turbiniformis was described and reported from Vene-
zuela only. Of the 241 species reported on in this paper, 39 have
been previously reported for the state. The remaining 202 species
are entirely new for the state.

ACKNOWLEDGMENTS

I wish to exj^ress my keen appreciation of the very helpful as-
sistance of Dr. A. J. Mix and of his encouragement and unfailing
interest throughout this work. I also wish to thank Dr. A. T.
Walker, Miss Mary Grant, and Mr. Maurice Hatch for the help
and special attention they gave to the writing of the Latin diagnoses
and Miss Edna Old for her timely help in the organization and
production of the plates.

Description of Species

CLASS MYXOPHYCEAE

Order CHROOCOCCALES

Family Chroococcaceae

Chroococcus limneticus Lemm.

(PI. I, fig. 3)

Cells paired, 4-32 in ;i colonial gelatinous mass. Individuals 8-16 mu in
diameter with the capsule, without the capsule 6-12 mu in diameter; pale
olive to light blue-green. The capsule is either distinct or obscured by
confluence in the colonial mass. Planktonic in temporary or permanent waters.

Coll. Sept., 1935, swamp in Chanute, 1935, 1936, 1937, Jackman's pond,
drainage ditch in Haskell bottoms, Lawrence.

Chroococcus turgidus (Kiitz.) Niig.

(PL I, fig. 1)
Cells generally single or 2-4, rarely as many or more than eight in a group.
Cell body blue-green or yellowish to brownish. Cells 13-40 mu in diameter
with the capsule, 8-32 mu without the capsule. Capsule colorless, generally
distinctly layered. Common, but never found in quantity, on mud bottoms
of still waters, but most frequently found among the filaments of green algae.

Thompson: Fresh-water Algae 9

Coll. Sept., Oct., 1935, Jackman's pond, Potter's pond. Haskell bottoms, boat
slough at north end of Ohio street, old quarry on East Fifteenth street,
Lawrence.

Chroococcus dispersus (v. Keiss) Lemm.

Cells in colonies of from 4-16 or more within a gelatinous envelope, the
cells either closely packed or widely dispersed throughout the mass, pale
to bright blue-green. Diameter with tiie ctipsule 5-6 mu, without the capsule 3-4
mu. The capsule is distinct, not layered and not confluent with the colonial
envelope, colorless. Planktonic in habit.

Coll. Sept. to Nov., 1935, and in the spring of 1936, Jackman's pond. City
park pool. East P'ifteenth street quarry, Lawrence.

Chroococcus caldarionnn Hansg.

(PI. I, fig. 2)

Aggregate cells forming a slimy mass, on wet soil, that is liver-red in color,
or bright blue-green when the red pigment is absent. This layer may cover
large areas. Cells purple-red to violet, 5-15 mu in diameter with the capsule,
3-6 mu in diameter without the capsule. Capsule more or less thick, not
layered and colorless. Common on moist soil and walls in greenhouses.

Coll. Jan. to May, 1935, 1936, 1937, Botany greenhouse, Kansas University,
Lawrence.

Gleocapsa mellea Kiitz.

(PI. I, fig. 5)

Aggregate cells form a crust on moist walls and moist earth which is pale-
flesh-colored to yellowish. Individual cells are 2.5-5.5 mu in diameter, spheri-
cal to angular, in globose to irregular colonies of two to many cells. Capsule
hyaline and colorless; cell body ficsh-colored to golden-yellow.

Coll. Dec, 1936, Chanute, on moist soil.

Gleocapsa magma (Breb.) Kiitz.

(PI. I, fig. 4)

Aggregate cells form a crumbly layer on moist cliffs which is copper-red
to purple-brown. Cells 6-12 mu in diameter with the capsule, without the
capsule 4.5-7 nui in diameter. Capsule layered, copper-red or colorless. Cell
body light blue-green.

Coll. Dec, 1936, on moist cliffs where water was seeping out, Chanute.

Gleocapsa aeruginosa (Carm.) Kiitz.

(PL I, fig. 6)

Aggregate cells form a crust, crumbly or slimj-, blue-green to gray-green.
Cells with the sheath 4-8.8 mu, without the sheath 2-3 mu in diameter, blue-
green. Few to many in a colony. Sheath colorless, rather thick, either not
at all or only slightly layered.

Coll. Dec, 1936, on a limestone cliff, Chanute.

Merismopedia punctata Meyen

(PI. I, fig. 8)

Colonies small, cells loosely aggregated, 2.5-3.5 mu in diameter, pale blue-
green. In standing water.

Coll. spring, 1936, Green's pond, Lawrence.

10 The University Science Bulletin

Merismopedia elegans A. Br.

(PI. I, fig. 9)

Colonies small or large, 16-4000 cells. Cells more or less close, 5-7 mu
wide, 5-9 long, light blue-green.

Coll. Oct., 1935, Jackman's pond, Lawrence.

Merismopedia elegans A. Br. ?

(PI. I. fig. 10)

Colonies small, cells large and closely aggregated, 10-15 mu wide, 7-15 mu
long. This species is being tentatively determined as M. elegans since its
dimensions overlap into those of M. elegans and even though the dimensions
are greater for the most part, the cell shapes are the same as those of
71/. elegans.

Coll. Sept.. 1937. Jackman's pond, Lawrence.

Merismopedia angularis sp. nov.

(PI. I, fig. 14)

Coloniae sunt satis grandes, crasso glutinoso involucro intectae. Cellulae
sunt 4.4 mu latae, 6.5 mu longae, 4-6 mu altae, pentagoniae, uno polo piano,
altero acuto. Cellulae in duplicibus ordinibus positae sunt. Plani poli cellu-
larum utriusque ordinis oppositi siml planis polls alterius ordinis. Acuti poli
apti sunt inter acutos polos proximi ordinis. Color corpori cellulae est clare
caeruleus. Involucrum coloniae est pellucidum.

Colonies moderately large with a thick gelatinous envelope. Cells 4.4 mu
wide, 6.5 mu long. 4-6 mu thick arranged in a flat plate with two rows of
paired cells alternating with two more rows of paired cells. Color of cell
body, a bright blue-green. Colonial envelope colorless.

Coll. July, 1937, Chanute,

Eucapsis alpina Clem, and Schantz
(PI. I, fig. 11)
Colony a cube of cells surrounded by a thick envelope. Cells a bright
blue-green. 7 mu long, 7.6 mu wide.

Coll. spring of 1937, Green's i)ond, Lawrence.

Aphanothece microscopica Nag.

(PI. I, fig. 12)

Colonial mass free-floating in standing water, small, ellipsoidal to amorphic,
V-j-2 mm. in size. Cells ellipsoidal to cylindric, single or in pairs, 4-4.5 mu wide,
lM;-2 times as long, bright blue-green. Among filaments of green algae in still
water.

Coll. Jan., 1936. City park pool. Jackman's pond, Lawrence.

Microcystis marginata (Menegh.) Kiitz.

(PI. I, fig. 7)
Colonies spherical or irregular, not composed of smaller division colonies
massed together. Cells spherical, 3.6 mu in diameter.
Coll. April, 1937, Green's pond. Lawrence.

Thompson: Fresh-water Algae 11

Order CHAMAESIPHONALES

Family Pleurocapsaceae

Pleurocapsa minor Hansg. em. Geitler

(PI. I, fig. 13)

Colonial mass consisting of a basal portion and an erect portion. The basal
portion forms a rhizoidal, branched system over the substrate. The erect
portion, also filamentous, is either uniseriate or multiseriate. Cells 3-9 mu in
diameter, pale blue-green.

Coll. Jan., 1935, Olathe, on Ba.vcladio chclonum.

Family Chamaesiphonaceae
Chamaesiphon cylindricus Boye

(PI. 1, fig. 15)

Sporangium cylindrical, without a definite foot, 2-2.5 mu wide. 11-13.2 mu
long, extension of the sheath thinner above than at the base. Endospores few,
generally 1-2. Epiphytic on filamentous green algae.

Coll. April, 1935. on Oedogonium and Rhizoclonium, boat slough at north
end of Ohio street, Lawrence.

Chamaesiphon gracilis Rhabh.

(PI. I, fig. 17)

Sporangium slightly stalked, gradually attenuated to a point from the base
outward, solitary or gregarious on the filaments of green algae. Sporangium
1.5-2 mu wide, 25-30 mu long.

Coll. Oct., 1935. on Cladophora. Jackman's pond. Lawrence.

Order HORMOGONALES
Suborder HOMOCYSTINEAE

Family Oscillatoriaceae

Spirulina major Kiitz.

(PI. I, fig. 20)

Trichomes 1-2 mu wide, non-septate, pale blue-green, winding 2.5-4 mu wide,
distance or lead of the turn 2.7-5 mu. In still water.
Coll. Oct., 1935, Jackman's pond. Lawrence.

Arthrospira jcnneri (Kiitz.) Stiz.

(PI. I, fig. 23)

Trichomes blue-green, not" at all or slightly constricted at the cros,s walls,
5-8 mu wide, the ends not attenuated. Winding 9-15 mu wide, lead 21-31 mu.
Cells short or almost square, and cell broadly rounded. In standing water.

Coll. Nov., 1935, June, 1937, Green's pond, Lawrence.

12 The University Science Bulletin

OscJllotoria gleophila Griin.

Trichomes 3.7-4.5 mu wide; cells as long as broad, cross wall more or less
constricted and more or less granulated. In the gelatinous material of Chaeto-
phora elegans.

Coll. March. 1936. East Fifteenth street quarry pool. Lawrence.

Oscillatoria curviceps Ag.

(PI. I, fig. 16)

Trichomes light or dark blue-green, slightly attenuated and curved at the
ends, not constricted at the cross walls, 10-17 mu wide. Cells one third to one
sixth times as long as broad, 2-5 mu long, cross wall much granulated. End-
cell not button-shaped, flat-rounded or truncate.

Coll. Dec, 1935, Jackman's pond, Lawrence.

Oscillatoria simplicissima Gom.

(PI. I, fig. 19)

Trichomes dark blue-green to yellowish blue-green, not constricted at the
cross walls, not attenuated at the ends, 8-9 mu wide. Cells one fourth to one
half as long as wide, 2-4 mu long, not granulated at the cross walls. End-cell
not button-shaped, half-round with or without a perceptible thickening of the
wall.

Coll. Jan., 1936, Potter's pond, University of Kansas campus, Lawrence.

Oscillatoria jenensis G. Schmid.

(PI. I, fig. 22)

Aggregate mass of trichomes dark brown. Trichomes 19.8-24.9. mu wide,
not constricted at the cross walls, gray-brown, somewhat spiraled at the ends.
Cells short, end-cell asymmetric-convex, not knobbed or thickened. Xo gran-
ulation at the cross walls.

Coll. Botany department greenhouse, 1935, Potter's pond, L'niversity of
Kansas campus, Lawrence.

Oscillatoria sancta Kiitz.

(PI. I, fig. 18)

Trichomes straight or bent, not distinctly constricted at the cross walls,
lightly attenuated at the ends, 10-20 mu wide, dark blue-green to dirty olive-
green. Cells one third to one sixth as long as broad, 2.5-6 mu long, not gran-
ulated at the cross walls. End-cell half-round and flattened, lightly knob-
like with a thicker cell wall on the end.

Coll. Sept.. 1935. Potter's pond, University of Kansas campus, Lawrence.

Oscillatoria gattulata van Goor.

(PI. I, fig. 21)

Trichomes bent, not constricted at the cross walls, two to four and one half,
mostly 3-4 mu wide, end-cell neither tapered nor knobbed. Cells almost
quadratic or up to twice as long as broad, 3^/-i-8 mostly 6 mu long. End-cell
broadly rounded, cross walls very obscure or indistinct. Cells very pale blue-
green with pseudovacuoles. There are fine granules present at the cross walls

Thompson: Fresh-water Algae 13

though not leguhirly enough to iiermit measurement of the cell length. The
C108S walls are most easily demonstrated by mounting the material in a
hanging drop over iodine crystals and allowing the drop to dry. The cover
slip is then mounted on a slide in water, the cross walls becoming quite
evident as the iodine diffuses out. though they do not show up while the
material is at first in the hanging drop.

Coll. March, 1937. drainage ditch in Haskell bottoms, Lawrence.

Phormidimn tenue (Menegh.) Gom.

(PI. I, fig. 24)

Trichomes straight or lightly- cur\-ed, very slightly constricted at the cro.ss
walls, attenuated at the ends, 1-2 mu wide, light, pale blue-green. Cells up
to three times as long as broad, 2.5-5 mu long, not granulated at the cross walls.

Coll. 1935. 1936, 1937, boat slough at north end of Ohio street, Jackman's
pond, Potter "s pond, common in cultures from ^-icinity of Lawrence.

Lyngbya Birgei Smith

(PI. I, fig. 25)

Trichomes straight, seldom curved, 20-24 mu wide. Sheath colorless, homog-
enous, 0.5-4 mu thick. Trichomes not constricted at the cross walls; end-
cell rounded but not attenuated. Cells shorter than broad. 2.5-5.5 mu long,
18-23 mu wide, many times with pseudovacuoles.

Coll. Dec. 1935, Potter's pond, Lawrence.

Lyngbya aerygineo-coerulea (Kiitz.) Gom.

(PI. I. fig. 27)

Filaments curved, sheath thin, colorless, not layered, the aggregate mass of
filaments dark blue-green. Cells 4-6 mu wide, 1-1% times as long, not con-
stricted at the cross walls, but sometimes granulated, pale blue-green. End-cell
truncated or rounded with a slightly thickened membrane.

Coll. Nov.. 1935. boat slough at north end of Ohio street, Lawrence.

Microcoleus vaginatus (Yauch.) Gom.

(PI. I, fig. 26)

Filaments single, creeping or joined into a greenish-black layer, wound and
much branched. Sheath colorless, thick. Cells 3.5-7 mu wide, 3-7 mu long,
often granulated at the cross walls and not constricted, bright blue-green.
End-cell knob-like.

Coll. 1937, Botany department greenhouse on moist soil, Lawrence.

Suborder HETEROCYSTINEAE

Family Nostocaceae
Aymbaena californica Borge.

(PI. II, fig. 4)

Trichomes straight and parallel with a hyaline sheath. Cells cylindrical,
slightly constricted at the cross walls, 5-5.5 mu wide. 5-6 mu long. Hetero-
cysts 6.5-7.5 mu wide, 10.5-12 mu long. Resting cells at some distance from

14 The University Science Bulletin

the heterocysts, one or two in a row, cylindrical, straight, rounded at the
ends, 6.5 mu wide. 14.5-17 mu long with a smooth outer sheath.
Coll. March, 1937, swamp, Chanute.

Nostoc muscorum Kiitz.

(PI. II, fig. 5)

Colony at first globular, later irregularly expanded over the substrate, lobed
by splitting and curled at the edges. Cells short-cylindric or orbicular, 3-4
mu wide, as long as broad. Resting cells oval, 4-8 mu wide, 8-12 mu long,
with a smooth yellow sheath. On moist earth and rocks and in greenhouses.

Coll. April, 1936, moist soil near Santa Fe pond, Chanute.

Nostoc fuscescens Fritsch var. mixta Fritsch

(PI. II, fig. 2)

Colonies globular with a .short stipe which attaches them to the substrate.
Colonial sheath yellow-brown. Trichomes curled and constricted at the cross
walls. Cells 3-4 mu wide, pale blue-green. Heterocysts oval to ellipsoidal,
5-7 mu wide, 6-7 mu long. Resting cells not known.

Coll. Dec, 1936, moist limestone cliff, Chanute.

Nostoc verrucosum Vauch.
(PI. II, fig. 1)
Colony at first spherical to ovoid, later becoming loosely flattened out over
the substrate. Colonial sheath thick and tough. The filaments at the pe-
riphery of the colony thickly intenvoven. Cells short-barrel-shaped, 3-3.5 mu
wide. Resting cells oval, 5 mu wide, 7 mu long with a smooth yellowi.sh outer
layer. Generally there is a fairly obvious radial arrangement of the filaments
from the center.

Coll. March, 1937, Jackman's pond, Lawrence.

Nostoc coeruleum Lyngb.

(PI. II, fig. 3)

Colonies spherical, free-living in standing water, with a thick, tough,
smooth sheath that is slightly if at all pale blue-green. Trichomes very thickly
massed. Cells short, keg-like in shape, 5-7 mu wide. Heterocysts broad-oval,
up to 10 mu wide. Resting cells unknown.

Coll. March, 1937, swamp in Chanute.

Cylindrospermum mains Kiitz.

(PI. II, fig. 6)

Layer dark olive-green. Cells cylindric or almost quadratic, 3-5 mu wide,
3-6 mu long, pale blue-green. Trichomes constricted at the cross walls.
Heterocysts long, somewhat broader than the vegetative cells, up to 10 mu
long. Resting cells single, ellipsoidal, 10-15.4 mu broad, 20-38 mu long, with
a brown papillose outer layer.

Coll. Oct., 1935, Botany department greenhouse, Lawrence.

Thompson: Fresh-water Algae 15

Cylindrospermum 7wmufi.ssnM!77n Collins

(PI. II, fig. 7)

Layer blue-green. Cells cj'lindrical, 2-2.7 mu wide, 4-7 mu long. Hetero-
cj'sts somewhat long, 4 mu wide, 6-8 mu long. Resting cells one or two in a
row, somewhat longish, 7-9 mu wide, 12-25 mu long, with a smooth, colorless
outer sheath.

Coll. April, 1936, Jackman's pond, Lawrence.

Family Scytonemataceae
Scytonema myochrous (Dillw.) Ag.

(PI. II, fig. 12)

Filaments joined in a dark-brown to dark-green layer on moist earth or
stones. Filaments 18-36 mu wide, 2-15 mm. long. False branching common,
either one or two branches coming from the same point on the main filament.
Cells 6-12 mu wide, quadratic or up to two times as long as broad, disc-shaped
on the ends of the trichomes. Heterocysts ovoid or rounded-quadratic.

Coll. Dec, 1936, on moist limestone cliffs, Chanute.

Plectonema Golenkiniana Gom.
(PI. II, fig. 11)
Filaments free, not forming a layer, intermingling with filamentous green
algae. False branching common with a single pseudobranch coming from a
point of origin on the main filament. Trichomes long, slender and surrounded
by a tough sheath that is colorless. Heterocysts are never present. Trichomes
1.2-1.8 mu wide.

Coll. Common in cultures from stagnant waters, 1935-1937.

Family Rivulariaceae

Calothrix Braunii Born, and Flah.

(PI. II, fig. 9)

Filaments straight, parallel, 9-10 mu wide, at the base weakly bulb-shaped.
Sheath colorless. Cells 6-7 mu wide, constricted at the cross walls, somewhat
shorter than broad. Heterocysts basal, hemispherical to half-ovoid.

Coll. Oct., 1935, Potter's pond, Jackman's pond, Lawrence.

Rivularia compacta Collins

(PI. II, fig. 10)

Trichomes with individual sheaths, sheaths confluent in the hemispherical
colony. Colony not lime encrusted. Trichomes with a basal heterocyst and
attenuated from the heterocyst outward.

Came up in culture, 1935, Lawrence.

Rivularia planctonica Elenk.

(PI. II, fig. 8)

Colony small. 0.5 mm. in diameter. Trichomes straight or curved, up to
300 mu long, 4.5-5.8 mu wide. Sheath narrow, colorless, scarcely visible.
Cells almost quadratic or somewhat longer than broad, with pseudovacuoles.
Heterocysts 8-9.8 mu long, single.

Came up in culture, 1935, Lawrence.

16 The University Science Bulletin

CLASS RHODOPHYCEAE
SUBCLASS PROTOFLORIDAE

Order BANGIALES

Porphyridivm cruentum Nag.

(PI. II, fig. 13)

Cells spherical, growing in a layer on damp soil, the layer appearing blood-
red. Resting cells contain a dark-red. stellate chloroplast with a central
pyrenoid.

Coll. Dec, 1935. 1936. damp soil, Chanute; 1936, damp soil in Botany
department greenhouse, Lawrence.

CLASS HETEROKONTAE

Order HETEROTRICHALES

Family Tribonemataceae

Tribonema bombycinum (Ag.) Derbes and Sol.

(PI. II, fig. 19)

Filaments consist of cells that are cylindrical or barrel-shaped, 2-5 times
as long as broad. Cells 5-20 mil wide (mostly between 7 and 13 mii). Cell
walls composed of H-shaped pieces which come apart readily. Chloroplasts
numerous and discoid. Reserve foods stored as oils or granules of leucosin.

Coll. 1936, on pots in the Botany department greenhouse, Lawrence.

Order HETEROSIPHONALES

Family Botridiaceae
Botrydium granulatum Wallroth

(PI. II, fig. 14)

Cells unicellular, multinucleate and terrestrial. The thallus consists of a
vesicular portion and a rhizoidal portion. The vesicular portion may be either
globose or dichotomously forked (if growing in a very shaded place). The
vesicular portion may be from 1-2 mm. in diameter, has a relatively tough
wall within which is a parietal sheet of cytoplasm with embedded chloro-
plasts. The material collected formed both hypnospores and aplanospores,
the whole mass assuming the shape of the original vesicle and having a
frosty appearance in contrast to the shining yellow-green of the vesicle.

Coll. 1936, damp soil. Botany department greenhouse, Lawrence.

Thompson : Fresh-water Algae 17

CLASS CHRYSOPHYCEAE
Order CHRYSOMONADALES
Suborder CHROMULININEAE

Family Chromulinaceae

Chromulina ovalis Klebs

(PI. II, fig. 17)

Cells solitaiy, free-swimming, miked, ovoid to spherical, pyriform or spindle-
shaped, with two parietal, plate-like chloroplasts, one on either side of the
protoplasts. Cells 6-7 nm wide, 9-14 mu long, fiagellum half again longer than
the body.

Coll. March, 1937, boat slough at north end of Ohio street, Lawrence. It
occurred in such quantity as to form a yellow-brown scum on the surface
of the water.

Family Mallomonadaceae
Mallomonas tonsurata E. Teil.

(PI. II, fig. 16)

Cells solitary, uniflagellate, with a firm oniamented periplast covered with
minute siliceous scales, from some of which arise long siliceous spines, ellip-
soidal, the posterior end rounded, the anterior end pointed. A single large
yellow chloroplast is parietal in position and appears as two from the side.
At the posterior pole are two contractile vacuoles. Cells 18 mu long, 7-8 mu
wide. The material collected was 15.2 mu long, 8.4. mu wide.

Coll. March. April. 1937, boat slough at north end of Ohio street, Lawrence.

Suborder ISOCHRYSIDINEAE
Family Synuraceae

Synura uvella Ehrenb.

(PI. II, fig. 15)

Cells radially united in spherical to oblong-ovoid colonies that are not
enclosed by a gelatinous sheath. The cells are pyriform with the anterior
end broadly rounded and the posterior end prolonged into a hyaline stalk.
The firm periplast is covered with siliceous scales that are arranged spirally
and the anterior end may be ornamented with short siliceous spines. There
are two laminate, curved chloroplasts arranged within the cell so that their
concave faces are opposed. There are two flagella of equal length at the
anterior end. Colonies 100-400 mu long. Cells 20-40 mu long, 8 to 17 mu wide.

Coll. March, April, 1937, boat slough at north end of Ohio street, Lawrence.

2—4141

18 The University Science Bulletin

Order CHRYSOTRICHALES

Family Phaeothamniaceae

Phaeothamnion Borzianum Pascher?

(PI. II, fig. 20)

In Febmary and March, 1937, the palmella stage of a species of Phae-
othamnion was collected growing on submerged stems in water coming from
a spring. The cells were loosely arranged in gelatinous tubes either in a
single row or in two rows in the tube with the cells alternating in position.
Single cells measured 7.6 mu in diameter and contained one to four parietal
chloroplasts which were more greenish-yellow than golden-brown. Although
accurate determination of Phaeothamnion species is not possible from the
palmella stage alone, this material may be tentatively determined as of the
species P. borzianum.

CLASS CRYPTOPHYCEAE

Order CRYPTOMONADALES

Family Cryptomonadaceae

Cryptomonas ovata Ehrenb.

(PI. II, fig. 18)

Cells solitary, free-swimming with two flagella of equal length. Cell body
long-ovoid, a little broader at the anterior end and obliquely truncate, often
weakly bent. A conspicuous gullet extends from the oblique anterior surface
almost two thirds of the length of the body. There are two large parietal
chloroplasts and may be numerous leucosin grains. Cells are either slightly
or greatly compressed, 20-80 mu long. 6-20 mu wide and 5-18 mu thick. The
material collected measvu'ed 16.5-19.8 mu wide, 39.6-46.2 mu long.

Coll. July, 1937, small temporary swamp and pool near Chetopa creek,
Altoona.

CLASS CHLOROPHYCEAE

Order VOLVOCALES

Family Chlamydomonadaceae

Chlamydomonas intermedia Chodat

(PL III, fig. 1)

Cells elliptical with a thin but clear membrane. The chloroplast is massive
and cup-shaped with one pyrenoid lying near the center of the cell. An eye-
spot is present near the anterior end. Cells biflagellate, 18-20 mu long.

Coll. March, 1936, Jackman's pond, Lawrence.

Chlamydomonas variabilis Dang.

(PI. Ill, fig. 6)

Cells broadly ellipsoidal to almost cylindrical with a very evident papilla
at the anterior end from which arise the two flagella. Flagella 1% times the
body length, 18-23 mu, width 12-15 mu. Eyespot at the side just above the
equator and protruding.

Coll. July, 1937, Village creek swamp, Chanute.

Thompson : Fresh-water Algae 19

Chlorogonium spirale Scherf. and Pasch.

(PI. HI, fig. 3)

Cells very narrow-spindle-shaped with a single, parietal, spiraled chloroplast.
The eyespot varies in position from the anterior end to the middle at the side.
One pyrenoid is present. Cells 20-40 mu long, generally only 3-5 mu wide.

Coll. July, 1937, Bun-ton. In a stock pool in the sand hills.

Chlorogonium euchlonim Ehrenb.

(PL III, fig. 2)

Cells broadly spindle-.shaped, the anterior pole drawn out into a snout
frequently, 25-70 mu long, 4-15 mu wide; the material collected measured
20-30 mu long, 8-13.2 mu wide. The chloroplast is diffuse and almost entirely
fills the cell. Pyrenoids are few to numerous, generally more than two, though
individuals with one and two and none at all were seen. Stages in asexual
reproduction and the formation of gametes were also seen in culture, though
no fusion of the gametes was seen.

Coll. Came up in an old culture from Santa Fe pond, Chanute.

Carteria cordijormis (Carter) Dill

(PI. Ill, fig. 5)

Cells very slightly compressed, oval-shaped in the narrow view, broadly
rounded in the wide view with a slight depression at the anterior end in which
the four flagella are inserted. The chloroplast is cup-shaped and maj' not
have a pyrenoid. An eyespot is evident, anterior in position.

Coll. March, 1937, Green's pond, Lawrence.

Family Phacotaceae
Phacotus lenticularis (Ehrenb.) Stein

(PI. Ill, fig. 7)

Solitary, free-swimming, biflagellate cells in which the protoplast lies withiij
an envelope composed of two overlapping halves. The outer portion of the
two halves is either smooth or sculptured and may be yellow-brown to very
dark colored and impregnated with calcium carbonate. The halves have a
slight flange on their opposed rims, are shallow so that they give a biconvex
shape to the cell in side and top view and a subcircular shape from the front.
Sexual reproduction has never been observed. In vegetative division the
gelatinous material between the protoplast and the halves increases in quantity,
pushing the halves apart, and the protopla.st divides to form four or eight
zoospores which may escape or remain within the old mother cell to form a
palmelloid stage. Cells 11.4-15.2 mu long, 7.6 mu wide.

Coll. March, 1937, swamp, Chanute.

Wislouchiella planctonica Skvortzow

(PI. Ill, fig. 4)

Cells strongly compressed; protoplast surrounded by a wall which has a
broad wing-like expansion and upon which are borne blunt cylindrical processes.
Front and side views show that there are two processes on each compressed
face: one process inserted at the cell's apex and projecting upward, the other
projecting downward from its insertion at the base of the protoplast. In

20 The University Science Bulletin

vertical view these processes are seen to be axial and opposite in position,
but the upper pair lies at an angle to the plane of the lower pair. The proto-
plast is ovoid to pyriform in front view and rhomboidal in vertical view.
Two long flagella are diagonally inserted. The chloroplast is massive and
cup-shaped; the eyespot lies some distance back from the anterior end. In
the material collected the cells measured 16.5-19.8 mu wide, 19-19.8 mu long;
the protoplast measured 13.2 mu wide, 132 mu thick and 16.5 mu long.
Coll. July, 1937, stock pond in the sand hills, Burrton.

Family Volvocaceae
Gonium sociale (Duj.) Warming

(PI. Ill, fig. 8)

Colony four-celled, arranged in the same plane with a matrical connection
between the individuals. Cells 10-22 mu long, 6-16 mu wide.

Coll. March, 1935, field ditch near Blue Mound, Douglas county, Lawrence.

In another collection. Green's pond, Lawrence, 1937, March, Gonium sociale
was found, in which the individuals were very loosely held together with a
seeming protoplasmic connection between the posterior poles. (PI. Ill, fig. 8b.)

Gonium jormosum Pasch.

(PI. III. fig. 9)

Colony sixteen-celled, cells farther apart than in G. pectorale with larger
spaces in between the connecting strands and no large matrix around the
border of the colony. Cells almost pyriform, with the base broadly rounded
to almost flat, 10.7 mu wide, 13.2 mu long. Chloropla.st large, cup-shaped and
containing a single pyrenoid.

Coll. July. 1937, Village creek, Chanute.

Pandorina morum (Miiller) Bory

(PI. Ill, fig. 10)

Colonies 4, 8, 16, or 32-celled, in the vegetative colony the cells closely
packed, in a reproducing colony cells wade apart, the colony Eudorina-Iike.
Cells pyriform or oblate-spherical, 9-17 mu long or wide. Reproducing colony,
PI. XIX, fig. 1.

Coll. June, 1937, drainage ditch in Haskell bottoms, Lawrence.

Eudorina elegans Ehrenb.

(PI. Ill, fig. 11)

Colonies spherical to obovoid, 16, 32, or 64-celled. Cells fairly wide apart,

16-24 mu long. Flagella two to four times the cell diameter; chloroplast

cup-shaped and containing several pyrenoids ; eyespot quite evident near the

anterior pole of the cell.

Coll. April, 1935, boat slough at north end of Ohio street, Lawrence.

Volvox perglobator Powers

Colony 1,000-1,500 mu in diameter with many thousand of equal-sized cells.
Dioecious, over a hundred egg cells occur in the posterior portion of the female

Thompson: Fresh-water Algae 21

colony, 50-100 antheridial cells in the male colony. Zygotes stellate with short
blunt warts on the outer wall.

Coll. June, 1937, temporary pond and swamp near Chetopa creek, Altoona.

Family Spondylomoraceae
S-pondyloTnorum quaternarium Stein

(PI. IV, fig. 1)

The colony consists of eight or sixteen loosely associated cells in alternating
tiers of four each and no colonial matrix. The anterior end of the cells is
broadly rounded and has a minute papilla on which are borne four flagella.
The posterior end is abruptly and definitely pointed. The protoplast is nearly
filled with a large chloroplast that may or may not contain a pyrenoid. The
eyespot is posterior in position. The colony is up to 50 mu long; cells 10-26
mu long, 8-15 mu wide.

Coll. March, 1935, came up in an infusion of hay and barnyard dirt, Law-
rence; July, 1937, temporary pond near Chetopa creek, Altoona.

Family Sphaerellaceae
Sphaerella lacustris (Girod.) Wittr.

(PI. IV, fig. 2)

The motile cells are ovoid to ellipsoidal, biflagellate and solitary. The
protoplast is somewhat pyriform and lies some distance in from the wall to
which it is connected by thin cytoplasmic strands. The fiagella lie in ge-
latinous canals which fork from the anterior pole of the protoplast. The
chloroplast contains several pyrenoids and may be obscured by the presence of
haematochrome which also obscures the eyespot that lies at the equator of
the cell. Zoospores up to 51 mu wide, up to 63 mu long; microgametes up
to 8 mu wide, up to 10 mu long; aplanospores 30-50 mu in diameter.

Coll. spring, 1936, Botany department greenhouse pool, Lawrence; Feb.,
1937, a rain barrel, Chanute.

Order TETRASPORALES

Family Tetrasporaceae

Tetraspora cylindrica (Wahelnb.) Ag.

(PI. IV, fig. .5)

Colony up to a meter long, 2-15 mm. thick. Cells in groups of four and
2-17 mu in length.

Coll. Feb., 1935, Olathe, in a small swift-running stream; March, 1937, Village
creek, Chanute.

Tetraspora gelatinosa (Vauch.) Desv.

(PI. IV, fig. 3)

Colony at first attached, later free-floating, up to 20 cm. thick, irregular in
shape, often held erect in standing water by the elaborated gas which collects
as a large bubble in the top of the sac. Cells 2-14 mu in diameter.

Coll. Feb., 1935, 1936, 1937, swamp near the boat slough at north end of
Ohio street, quarry pool on East Fifteenth street, Lawrence.

22 The University Science Bulletin

Family Coccomyxaceae
Ourococcus bicaudatus Grob.

(PL IV, fig. 4)

Cells very variable in form from egg-shaped to spindle-shaped, with either
both ends sharply drawn out into a point or one end rounded and the other
pointed and either slightly curved or greatly bent. Cells 4-9 mu wide, 2-4 times
as long. Chloroplast parietal with one pyrenoid.

Coll. July, 1937, temporary pond in the sand hills, Burrton.

Coccomyxa dispar Schmidle

(PI. IV, fig. 6)

Cells cylindrical and irregularly placed throughout a colonial matrix which
is made up of the confluent sheaths of the individuals. The chloroplast is a
longitudinal, parietal plate which does not encircle the cell and which lacks
a pyrenoid. Cells 2.5-4 mu wide, 6.5-10 mu long. Those collected measured
2.5 mu wide and 5.8 mu long.

Coll. March, 1936, East Fifteenth street quany pool, Lawrence.

Nannochloris hacillans Naumann

(PI. IV, fig. 7)

Cells solitary and without a gelatinous envelope; subspherical to sub-
cylindrical in shape. There is one discoid chloroplast without a pyrenoid at
one end of the cell. Cells less than 4.5 mu long, those collected measured 3.8
mu long, 3 mu wide.

Coll. Jan., 1936, Jackman's pond, Lawrence.

Chlorosarcina consociata (Klebs) G. M. Smith

(PL IV, fig. 8)
Cells in flat packets of four to twenty with a distinct envelope around
the plant mass. Chloroplasts parietal with a single pyrenoid. Cells 18-25 mu
wide. Those collected were 19.6 mu in width.

Coll. March, 1936, swamp near boat slough at the north end of Ohio street,
Lawrence.

Order ULOTRICHALES

Suborder ULOTRICHINEAE

Family Ulotrichaceae

Ulothrix tenuissima Klitz.

(PL IV, fig. 9)

Cells 15-19.4 mu wide. In j^oung filaments the cells may be one fourth as
long as wide. Chloroplast a wide band with two or more pyrenoids that does
not completely encircle the cell.

Coll, Feb., 1935. 1936, Kaw river, Lawrence; 1937, Village creek, Chanute.

Thompson: Fresh-water Algae 23

SticJwcoccus subtilis (Ktitz.) Kleckcr

(PI. IV, fig. 12)

Cells cylindrical, with a parietal, laminate chloroplast that does not encircle
more than half of the cell and which contains one pyrenoid. The cells may
be solitary, in short filaments of a few cells or longer filaments of several
cells. The cells vary from 5-9 mu in length and average 7 mu in width.
Solitary cells and the end cells of filaments have the outer end wall rounded.

Coll. March, 1935, 1936, 1937, Kaw river, Lawrence; April, 1935, 1937,
swamp, Chanute.

Gleotila 'protogenita Kiitz.

(PI. IV, fig. 10)

Cells 3.5-4 mu thick, 1-2 times as long. Chloroplast large, as long as the
cell, parietal and almost completely encircling the cell, without pyrenoids.
The filaments are gradually constricted at the cross walls.

Coll. March, 1935, Jackman's pond, Lawrence.

Geminella ordinata West and G. S. West

(PI. IV, fig. 15)

Filaments composed of cells equidistant from one another with a gelati-
nous tube; cells 5.8 mu thick. The individual cells are broadly rounded at
the poles and contain a parietal, laminate chloroplast, having one pyrenoid,
which encircles more than half of the cell.

Coll. April, 1936, Lawrence.

Hormidiuni jiaccidum A. Br.

(PI. IV, fig. 11)

Cells 12 mu thick, IV2 to 2 times as long and arranged in long filaments
in standing water. Chloroplast parietal and nearly filling the length of the
cell. Pyrenoids are scarcely discernible.

Coll. March, 1935, 1937, boat slough at north end of Ohio street, Lawrence.
April, 1935, 1937, swamp, Chanute.

Family Microsporaceae
Microspora stagnorwn (Kiitz.)

(PI. IV, fig. 14)

Filaments not constricted at the cross walls. Cells from 5 to 9.5 mu in
width and from 1 to 4 times as long. Chloroplast parietal, covering both
the ends and sides of the cell. It is an irregularis^ expanded sheet, perforate
and reticulate and lacks pyrenoids.

Coll. Jan., 1936, from a field ditch near Blue Mound, Lawrence.

Microspora tumidula Hazen

(PI. IV, fig. 13)

Cells almost cylindric, somewhat thinner at the cross walls, 6.7 to 9.5 mu
wide and 1 to 2 times as long. Chloroplast somewhat thicker than that of
the above species and covering a larger portion of the cell wall.

Coll. Dec, 1935, ditch on Ea.'^t Fifteenth street, Lawrence.

24 The University Science Bulletin

Family Cylindrocapsaceae
Cylindrocapsa geminella WoUe

(PL IV, fig. 16)

Cells in a single series within a thick gelatinous tube, or rarely irregularly
arranged side by side within the same tube, each cell surrounded by several
concentric strata. Width of the cells varies from 10 to 18 mu. The chloroplast
is massive and contains a single pyrenoid which is generally obscured by nu-
merous starch grains..

Coll. April, 1935, swamp, Chanute.

Family Chaetophoraceae
Stigeoclonium tenue (Ag.) Kiitz.

Cells united to form a branching filament which usually grows attached to
submerged objects. A prostrate portion which attaches the alga to the sub-
strate is pseudoparenchymatous or irregularly branched and gives rise to
numerous erect filaments which also branch. The ends of the branches are
attenuated to sharp points and the branching is opposite. Cells 7 to 15 mu
wide and 1 to 3 times as long; cylindrical or slightly swollen in the middle.
The cells of the main axis which give rise to side branches are generally 10 mu
long and 10 mu wide in the older portion of the filament. The chloroplasts of
the older cells are transversely zonate and contain many pyrenoids; those of
the smaller, younger cells encircle the length of the cell and contain a single
pjTenoid.

Coll. April, May, 1935, 1936, 1937, Kaw river, Lawrence; 1935, 1936, swamp,
Chanute.

Chaetophora incrassata (Huds) Hazen

(PI. IV, fig. 19)

Colony an irregular elongate mass of intertwined filaments forming an axial
strand from which arise numerous groups of short lateral branches, the
whole embedded in a copious jelly and macroscopic in size. The ends of
some of the branchlets taper out into long hyaline hairs, ending in a point.
The chloroplast of the older, larger cells is a transversely zonate band with
several pyrenoids; that of the younger cells fills the entire cell and contains
one pyrenoid.

Coll. April, 1935, 1937, swamp, Chanute; 1935, Deer creek. Garnett.

Chaetophora clegans, (Roth) Ag.

(PI. IV, fig. 17)

Colony macroscopic, globular or hemispherical, depending on whether it is
growing attached to submerged twigs or stones. The branches radiate through-
out the gelatinous matrix from the point of attachment of the colony. The
branching at the ends of the main filaments is fasciculate, but further within
the colony they arise at widely separated points on the main filament. Cells
of the main filament are 6 to 15 mu thick and 3 to 10 times as long.

Coll. March, April, May, 1935, 1936, 1937, Jackman's pond. East Fifteenth
street quarry pool, Lawrence.

Thompson: Fresh-water Algae 25

Draparnaldia plumosa (Vauch.) Ag.

(PI. IV, fig. 20)

The plant body is macroscopic and divided into two types of branches.
The primary branches are made up of large cylindrical or barrel-shaped cells
50 to 90 mu thick; while the lateral branches, which are 6 to 10 mu thick,
grow in fasciculate groups and have the terminal cells of their filaments drawn
out into long hyaline setae. The length of the cells of the primary filament
is from 1 to 3 times the width and all the cells of the filament are about the
same length. The chloroplast of the primary cells may be entire or reticulate,
with smooth to toothed edges, and always contains several pyrenoids. The
chloroplast of the lateral branch cells covers the entire wall surface of a cell
and generally contains a single pyrenoid.

Coll. April, 1935. 1937, boat slough at north end of Ohio street, Lawrence;
1936, swamp, Chanute.

Draparnaldia Ravenelii Wolle

(PI. IV, fig. 18)

The general description of this species is nearly the same as that of the
foregoing species, but the two differ greatly in the general shape of the fascicle
of lateral branches. Those of D. plumosa are lanceolate-elongate in outline,
while those of D. ravenelii are orbicular in outline and much more densely
branched. Also, the cells of the main filament of D. plumosa are generally
longer than broad, while the cells of the main axis of D. ravenelii are shorter
than broad to cuboidal.

Coll. April, 1935, roadside ditch, Labette.

Microthamnion Kuetzingianum Nag.

(PI. V, fig. 1)

The plant body of this alga is densely branched with all the cells of the
thallus of the same diameter. The branches originate as lateral outgrowths
from the upper end of a cell and there is a marked tendency for the branches
to be unilateral. There is a single, parietal, laminate chloroplast extending
the whole length of the cell and partially encircling it which does not con-
tain pyrenoids.

Coll. April, May, 1935, boat slough at north end of Ohio street, Lawrence.

Ulvella involvens (Savi) Schmicile

(PI. V, fig. 3)

The thallus of this alga is composed of a sessile, orbicular pseudoparenchy-
matous layer of cells that grows epiphytically on other plants, animals or
other objects. Young thalli are one cell in thickness, but older thalli may be
several cells in thickness in the central portion. The cells are multinucleate
and contain a single parietal, reticulate chloroplast with or without a pyrenoid.

Coll. March, 1936, on a turtle, Olathe.

Chaetopeltis orbicularis Berth

(PI. V, fig. 6)
The thallus of this alga is discoid, usually one cell layer in thickness, and
grows epiphytically on the filaments of other algae. The thallus has a thin

26 The University Science, Bulletin

gelatinous sheath from which arise numerous long, delicate, gelatinous setae. '
The cells contain a single, parietal, laminate chloroplast which covers most of
the cell's free face and has a single pyrenoid.

Coll. April, May, 1935, 1936, on Cladophora glomerata in Kaw river, Law-
rence.

Aphanochaete repens A. Br.

(PI. V, fig. 2)

The filaments of this alga grow epiphytically on the filaments of the larger
algae and consist of simple or irregularly branched short filaments which taper
at their extremities. The cells are globose to globose-cylindrical or barrel-
shaped. Those in the middle of a filament have a single parietal, laminate
chloroplast usually containing many pjnenoids. The chloroplasts of the cells
at the extremities of the filaments contain only one pyrenoid. From the
dorsal side of the cells one or more slender, hyaline, unicellular hairs may
arise, tapering from a bulbous base to a fine point, and are probably modified
branches.

Coll. March, April, 1935, 1936, on Cladophora sp., Kaw river, Lawrence.

Family Protococcaceae
Protococcus viridis Ag.

The cells of this alga grow either singly or in packets of two, four, or more.
The solitary cells are spherical or ellipsoidal with a thick wall that is without
a gelatinous envelope. The cells contain a single, parietal, laminate, chloro-
plast that is usually without pyrenoid, but may have one. It is commonly
found growing epiphytically on the leeward side of trees from the prevailing
wind, or on damp walls or surfaces.

Coll. throughout the years 1935, 1936, 1937, Lawrence.

Family Coleochaetaceae
Coleochaete scutata Breb.

(PL V, fig. 4)

The thallus of this alga is an orbicular parenchymatous layer of cells, one
cell laj^er in thickness, from certain cells of which arise single, long unbranched,
cytoplasmic setae. The cells are angular by mutual compression and contain
a single, parietal, laminate, chloroplast that partially or wholly encircles the
protoplast and usually contains one large pyrenoid.

Came up in culture, 1936, Lawrence.

Suborder CLADOPHORINAE

Family Cladophoraceae

Cladophora glomerata (L.) Kiitz.

(Pl. V, fig. 7)

This alga forms macroscopic, branching thalli which consist of large, mu\-

tinucleate cells, five to twenty times as long as broad, and which are united

end to end to form branching filaments that appear subdichotomous, but

which are generally lateral in branching. The branches originate at the upper

Thompson: Fresh-water Algae 27

end of a cell and are opposite. The thalli are usually sessile and grow attached
to submerged objects. The cells of the main filament are 130 mu thick and
those of the small branches are twenty-two mu thick at the least. A single,
large, parietal, reticulate chloroplast covers the entire inner surface of the
cell wall and contains numerous pyrenoids.

Coll. throughout the years 1935, 1936, 1937, Kaw river, Lawrence.

Rhizoclonium hieroglyphicum (Ag.) Kiitz.

Cells 10 to 40 mu thick and two to eight times as long as broad; united to
form long, unbranched filaments that are fastened to the substrate by uni-
cellular, rhizoidal branches at the bases. Chloroplast a parietal, reticulate
sheet that covers the entire cell wall and contains numerous pyrenoids which
are placed at certain intersections of the reticulum. The cells are multinu-
cleate.

Coll. 1935. swamp, Chanute; 1936, Jackman's pond. Potter's pond, Lawrence.

Rhizoclonium crispum Kiitz.

(PI. V, fig. 5)

The cells of this species are never more than twice their breadth in length.
They are mostly 20 to 50 mu thick, very irregular in shape and have a
membrane which is 3 to 4 mu thick. The filaments tend to grow in tangled
masses and may be collected in such masses among other free-floating algae.

Coll. May, 1936, boat slough at north end of Ohio street, Lawrence.

Basidadia chelonum (Collins) Hoffm. and Tilden

(PI. V, fig. 9)

This alga has been found growing on the carapace of turtles only, to which
it is attached by rhizoidal outgrowth at the end of an elongated basal cell.
The branching is from the ba.se only or rather very rarely from a cell higher
up on the erect filament, and the cross wall of the cell is generally a bit
remote from the main filament. The cells are 35 mu thick or less and the
chloroplast is like that of CJadophora.

Coll. fall, 1935, Newton; April, 1935, Olathe; July, 1937, Newton.

Pithophora kewensis Wittr.

(PI. V, fig. 8)

This alga resembles Cladophora, but is markedly different by the presence of
very dark terminal and intercalary akinetes, which are thicker-walled than
the vegetative cells. In culture it grows most luxuriantly in the darker shaded
portion of the container. The akinetes are visible to the unaided eye and
give the filaments a beaded appearance. The filaments are less than 150 mu
thick, those of the material determined being 105 mu thick and the akinetes
being up to 140 mu thick. The branches, according to the published descrip-
tions, originate just below the upper distal portion of a cell; however, in the
material determined, the branches were, for the most part, arising from the
mid portion of the cells, though they varied in this. Also, there were occasion-
ally opposite branches.

Coll. 1935. Merrit Lake. Fort Leavenworth.

28 The University Science Bulletin

Pithophora varia Wille

Cells less than 100 mu thick; akinetes very variable in shape; otherwise
the species is quite like P. kewensis.
Coll. July, 1937, Neodesha.

Order OEDOGONIALES

Family Oedogoniaceae

Oedogonium crispum (Hass.) Wittr.

(PI. V, fig. 11)

Oogonium single, egg-shaped, opening by a circular slit around the upper
part to release the oospore. Oospore wall smooth. Vegetative cells 12-16 mu
thick and generally from 3-4% times as long as broad. Oogonium 37-43 mu
thick, antheridial cells 8-14 mu thick and 7-12 mu long.

Coll. April, 1936, boat slough at north end of Ohio street, Lawrence.

Oedogonium glohosum Nordst.

(PI. V, fig. 10)

Oogonium single, subspherical ; pore in the upper part. Oospore subspheri-
cal, almost filling the oogonium; with a smooth membrane. Antheridium
1-7 celled. End-cell bristle-like. Vegetative cells 10-14 mu thick, 4-7 times
as long. Oogonium 32-40 mu wide, 32-46 mu long; oospores 30-37 mu wide,
28-37 mu long. Antheridial cells 9-12 mu wide, 4.5-8 mu long.

Coll. April, 1937, Green's pond, Lawrence.

Oedogonium cyathigenmi Wittr. sec. Hirn

(PI. VI, fig. 1)

Oogonium single, or 2-3 in a row, egg-shaped to quadratic-ellipsoidal with a
pore in the upper part. Oospore the same shape as the oogonium and filling
it. Epispore smooth, mesospore longitudinally ribbed, the ribs distinct but
sometimes anastomosed. Androsporangium many-celled. Dwarf males cup-
shaped with an internal antheridium, bent, generally attached to the basal cell
beneath an oogonium rather than to the oogonium. Vegetative cells 21-30
mu thick, 2-10 times as long. Cell just beneath the oogonium 42-48 mu wide,
up to two times as long; oogonium 57-66 mu wide, 70-100 mu long. Oospore
51-62 mu wide, 60-75 mu long. Androsporangial cell 23-30 mu wide, 12-30
mu long. Dwarf male 12-15 mu wide, 50-58 mu long.

Coll. June, 1937, Chetopa creek, Altoona.

Bulbochaete varians Wittr. sec. Hirn

(PI. VI, fig. 2)

Gynandrosporous. Oogonium ellipsoidal, terminal or beneath a terminal
hair, or under an androsporangium or under a vegetative cell which bears an
androsporangium. Androsporangium elongate, epigynous or nearly so, 1-2
celled. Dwarf males with an outer 1-3 celled antheridium, attached near to
or on the oogonium. Vegetative cells 17-22 mu thick 1^/4-1Mj times as long.
Oogonium 30-36 mu wide, 44-54 mu long. Androsporangial cells 14-17 mu
wide, 14-18 mu long. Stalk of the dwarf male 14-16 mu thick, 24-27 mu long.
Antheridial cells 8-10 mu wide, 6-7 mu long.

Coll. June, 1937, Green's pond, Lawrence.

Thompson: Fresh-water Algae 29

Order ULVALES

Family Schizomeridaceae

Schizomeris Leibleinii Kiitz.

(PI. VL fig. 3)

Filaments of this alga consist of an elongate basal cell with a discoid hold-
fast, a region in which cell division has been in one plane and a region in
which cell division has been in two planes so as to make a solid cylinder of
cells. The chloroplast in the basal cell and first region is a ulotrichoid type
and encircles about two thirds of the protoplast. They usually contain
several pyrenoids. The chloroplast of the cells in the cylindrical portion of
the thallus is more massive and fills most of the protoplast. The apical
cell of a thallus is acute. Quadrifiagellate zoospores are formed in either
terminal or intercalary sporangia, but always in the upper portion of the
thallus. The thick lateral walls of the simple filament persist as rings after the
vertical division of the cells and their multiplication to form the solid cy-
lindrical thallus.

Coll. June, 1937, drainage ditch in Haskell bottoms, Lawrence.

Order SCHIZOGONIALES

Family Schizogoniaceae
Schizogonium murale Kiitz.

(PI. VI, fig. 6)

Thallus a simple, cylindrical filament of cells within a sheath. Cells wider
than long, 7-7.5 mu wide, 5-5.5 mu long. Thej' contain a single axial, stellate
chloroplast which has one pyrenoid. •

Coll. April, 1937, Green's pond, Lawrence.

Order CHLOROCOCCALES

Family Chlorococcaceae

Chlorococcum infusionum (Schrank) Menegh.

(PI. VI, fig. 5)

Cells spherical; either solitary or aggregated in a gelatinous mass, 10-15 mu
in diameter. The chloroplast of young cells is a massive cup and contains one
pyrenoid; in older cells it becomes diffuse and contains several pyrenoids.
The cell is at first uninucleate, but contains many nuclei just before reproduc-
tion. Reproduction is by zoospores.

Coll. April, May, 1935, 1937, field ditch near Blue Mound, Lawrence.

Chlorococcum humicola (Nag.) Rab.

(PI. VI, fig. 4)

Cells very variable in size, 2-3 mu seldom up to 25 mu in diameter, single
or in small groups. A single parietal chloroplast which contains one pyrenoid
is present.

Coll. Dec, March, 1935, 1936, on damp soil in Botany department green-
house.

30 The University Science Bulletin

Golenkinia radiata Chod.

(PL VI, fig. 8)

Cells solitaiy, spherical, thin-walled, the wall bearing many long, delicate
setae that are not thickened in the portion next to the wall. Chloroplast
cup-shaped and containing one pyrenoid. Reproduction bj' autospores.

Coll. March, April, 1937, small aquarium, Lawrence.

Family Endosphaeraceae
Chlorochytrium lemnae Cohn

(PI. VI, fig. 7)

Cells endophytic in the tissues of species of Duckweed or Lemna, irregularly
globose to ellipsoidal, with relatively thin, homogenous cell walls. Chloro-
plast at first parietal and cup-shaped, later becoming radially vacuolate and
filling the entire cell lumen.

Coll. May. 1935, growing in Lemna minima, Lawrence.

Family Characiaceae
Characium ornithocephalum A. Braun

(PI. VI, fig. 9)

This alga may be epiphytic on other algae, submerged sticks, stones or on
aquatic animals. The material collected was growing on Oedogonium. The
cells are roughh' half-moon-shaped or as the specific name suggests, they
are somewhat the shape of a bird head. Cells without stalk are 25-33 mu
long and generally half as thick. The cells are vniinucleate and have a
parietal laminate chloroplast which contains one pyrenoid. The material
collected has been placed tentatively as this species since there was no other
that had the same peculiar cell shape and the measurements were approxi-
mately the same. The material determined varies from 15-30 mu in length
and 5-10 mu in width. The stalk of the collected material was very short
compared to the stalk figured in Pascher for C. ornithoce-phaluni.

Coll. Ma3% 1935, on Oedogonium sp.. boat slough at north end of Ohio
street, Lawrence.

Characium angustum A. Braun

(PI. VI, fig. 10)

Cells 22.5 mu long, 7.5-9 mu wide. The material determined as of this
species fits the description but not the figure in Pascher for this species. In
culture a dense growth was obtained on the wall of the container from which
pure mounts could be made. The disk at the end of the stipe was very
distinct and the rounded base of the cell where it changed abruptly into stipe
was also clear. This feature in itself does not fit any of the described species
in Pascher, nor any of the figures.

Coll. April, May, 1935, on Oedogonium sp., Jackman's pond, Lawrence.

Characium Naegelii A. Braun

(PI. VI, fig. 11)

Young cells linear-lanceolate; older cells almost elliptic, with a short stipe
that is not broadened into a disk holdfast at the base. Cells 20-42 mu long.
7-18 mu wide; stipe 4 mu long.

Coll. spring, 1936, boat slough at north end of Ohio street, Lawrence.

Thompson: Fresh-water Algae 31

Characium Braunii Breug.

(PI. VI, fig. 12)

Cells 25-32 mu long, 6.5-13 mu wide with a short stipe which is a quarter to
a fifth as long as the cell body. The cell body is equally attenuated at each
end, linear-lanceolate. The basal disk may be colored brown.

Coll. spring, 1937, on Cladophora, Kaw river dam, Lawrence.

Family Hydrodictyaceae
Pediastrum Boryanum (Turp.) Menegh.

(PI. VI, fig. 13)

Cells united to form a coenobium one cell layer in thickness, without
any perforations or space between calls. The coenobia may contain 4-128
cells. The marginal cells have two projections and the cells in general
measure up to 40 mu in diameter.

Coll. April, May, 1935, 1936, Jackman's pond, quarry pool on East Fifteenth
street, Lawrence; Feb., 1935, Newton.

Pediastrum Boryanum (Turp.) Menegh. var. brevicorne

Al. Braun

(PI. VI, fig. 14)

Coenobium with 8-32 cells. Cells up to 35 mu wide. Border cells rounded
out. Processes short, 4 mu long, 2-3 mu wide. Two forms occur; smooth
(forma glabra), punctate (forma punctata).

Coll. April, 1936, Jackman's pond, Lawrence.

Pediastrum Boryanum (Turp.) Menegh. var. longicome Reinsch

(PI. VI, fg. ly)
Coenobium eight or more celled, the inner cells 5-6 sided. The marginal
cells have two abrupth^ projecting, slender processes from each cell.

Coll. April, 1937, swamp by the boat slough at the north end of Ohio street,
Lawrence.

Pediastrum duplex Meyen var. cornutum Raciborski

(PI. VI, fig. 15)

Number of cells in the coenobia variable, 4-8-16, etc. The processes of the
marginal cells are very abrupt and smooth with blunt tips.
Coll. May, 1935, Jackman's pond, Lawrence.

Pediastrum duplex Meyen var. reticulatum Lager.

(PI. VI, fig. 10)

Coenobium 8-16-celled, the cells almo.<t H-shaped, perforations very large;
marginal cells 12-18 mu in diameter.
Coll. spring, 1936, Lawrence.

Pediastrum duplex Meyen var. gracillimum W. and G. S. West

(PI. VI, fig. 18)

Coenobium up to 87 mu in diameter. Marginal cells very deeply incised by
the presence of two long, slender, bhmt-tipped processes from each cell. The
interior cells of the coenobiimi are four-armed.

Coll. spring, 1936, field ditch near Blue Mound, Lawrence.

32 The University Science Bulletin

Pediastrum simplex (Meyen p. p.) Lemm. var. radians Lemm.

(PI. Vll, fig. 3)

Marginal cells with one process; center of the coenobium one large opening
so that the coenobium is merely a ring of marginal cells.
Coll. April, 1935, Jackman's pond, Lawrence.

Pediastrujn clathratum (Schroeter) Lemm.

(PI. VII, fig. 2)

Coenobium with large and small perforations. The middle cells many-sided
and the marginal cells forming an oviter ring, each cell of which has a long
process which is slightly concave at the tip.

Coll. spring, 1935, quarry pool on East Fifteenth street, Lawrence.

Pediastrum clathratum (Schroeter) Lemm. var. duodenarium

(Bailey) Lemm.

(PI. VII, fig. 4)
Coenobium 16-celled with five cells in the middle and the remainder forming
a ring about them.

Coll. spring, 1935. Jackman's pond, Lawrence.

Pediastrum duplex Meyen var. clathratum A. Braun

(PI. VII, fig. 1)

-Coenobium 8-64-celled. Cells 20 mu wide, 25 mu long. Middle cells quite
different from the marginal cells, their walls being rounded out. Perforations
quite large.

Coll. June, 1937, Moulton's lake, Neodesha.

Pediastrum tetras (Ehrenb.) Ralfs

(PI. VI, fig. 17)

Coenobivmi 4 to 16-celled. Marginal cells with a narrow sinus which reaches
to the middle of the cell. Middle cells many-sided, each with a narrow slit
on one side.

Coll. May, 1937, boat slough at north end of Ohio street, Lawrence.

Hydrodictyon reticulatimi (L.) Lagerh.

Coenobium made up of cells united in such a fashion as to make a closed
net in which the meshes are generally six-sided. The cells are at fii'st uninucle-
ate, but become multinucleate as growth in size continues. The chloroplast is
at first zonate and contains one pyrenoid; later in the older cells it becomes
reticulate with many pj^renoids. Reproduction either by biflagellate zoospores
or biflagellate gametes.

Coll. spring, 1935. 1936, Merrit Lake, Fort Leavenworth.

Family Coel.\straceae
Coelastrum reticulatum G. S. West

(PI. VII, fig. 7)
Coenobium spherical, composed of tetrad-groups of cells, the groups united
by long processes leaving large spaces in between. The cells of a group of four

Thompson: Fresh-water Algae 33

are closely ]iacked witli little if anj' space between iudixidual cells. Cells
6-10 mu in diameter.

Coll. June, 1937, boat slough at north end of Ohio street, Lawrence.

Coelastrum microporum Nag.

(PI. VII, fig. 6)

Coenobium a hollow sphere of 4, 8, 16, 32 up to 128 cells. Cells spherical,
6-27 mu in diameter, walls smooth, united by short processes. Chloroplast
of young cells cup-shaped with one pyrenoid, later in older cells becoming
diffuse and filling the entire cell cavitj'.

Coll. spring, 1935, boat slough at north end of Ohio street, Lawrence; June,
1937, Chetopa creek, Altoona.

Coelastrum sphaericum Nag.

(PI. VII, fig. 5)

Cells more ovoid than spherical, closely appressed, the space between the
cells being at the most one half the diameter of the cells. Cells up to 25 mu
in diameter.

Coll. spring, 1935, boat slough at north end of Ohio street, Lawrence; June,
1937, Chetopa creek, Altoona.

Family Oocystaceae

Chlorella conductrix Brandt

(PI. VII, fig. 8)

Cells solitaiy, endophytic in protozoa, spherical. They are commonly found
living within the cells of Hydra, Stentor and Paramecium.

Coll. 1935, 1936, 1937, in Paramecium and Chilodon, Potter's pond, Jack-
man's pond, Lawrence.

Westella botryoides (W. West) de Wildm.

(PI. VII, fig. 9)

Cells spherical to subspherical. quadrately grouped in fours or eights in
irregular free-floating colonies containing a variable number of cells up to
lOO. The groups of cells are held together by old mother-cell wall remains.
Cells 3.5-8 mu in diameter. Chloroplasts parietal and cup-shaped or entirely
filling the cell. There may or may not be a pyrenoid.

Coll. June, 1937, drainage ditch in Ha.skell bottoms, Lawrence.

Dictyosphaerium pulchellum Wood

(PI. VII, fig. 10)

Cells spherical, united by dichotomously forked threads, the persisting
mother-cell walls of previous generations. The whole is embedded in a
copious, colorless, gelatinous sheath. Chloroplast parietal and cup-shaped with
one pyrenoid. Cells 5-9 mu in diameter.

Coll. March, 1936, Lawrence. In Botany greenhouse pool.

3^141

?4 The University Science Bulletin

Trochisda pachy derma (Reinsch) Hansg.

(PI. VII, fig. 11)
Cells solitary, spherical, 12-19 mil in diameter. Inner layer of the cell wall
strongly warty with 7-20 warts with blunt outer surfaces.

Coll. June, 1937, Moulton's lake, Verdigris river, Neodesha.

Excentrosphaera viridis G. T. Moore

(PI. VII, fig. 12)

Cells ellipsoidal to subspherical, 22-55 mu in diameter, with numerous an-
gular chloroplasts that are peripheral in arrangement. The cell wall is irregu-
larly thickened and often stratified in the thickened portion. Reproduction
is by the formation of aplanospores which are released through a pore in the
cell wall.

Coll. July, 1937, Green's pond, Lawrence.

Oocystis parva W. an(i G. S. West

(PL VII, fig. 14)

Cells 2, 4, or 8, surrounded by the partially gelatinized and greatly expanded
mother-cell wall. Individual cells 6-12 mu long, 4-7 mu wide; rather narrowly
elliptic with the poles pointed, but without nodular thickenings. Chloroplasts
2 or 3 and parietal, with or without pyrenoids. Cells may be solitary by escape
from the mother-cell wall.

Coll. April, 1936, Green's pond, Jackman's pond, Lawrence.

Oocystis elliptica W. West

(PI. VII, fig. 15)

Cells elongate-elliptic, without thickened poles, 24-25 mu long, 11-11.5 mu
wide, solitary or in four- or eight-celled aggregates within the old mother-cell
wall.

Coll. July, 1937, drainage ditch in Haskell bottoms, Lawrence.

Dimorpho COCCUS lunatiis A. Br.

(PL VII, fig. 16)

Cells arranged in tetrads, the tetrads united to one another in irregular free-
floating colonies by the branching remains of old mother-cell wall. Cells
oval to elliptic, 10-20 mu long.

Coll. July, 1937, Green's pond, drainage ditch in Haskell bottoms, Lawrence.

Closteridium lumda (Reinsch) Wille ?

(PI. VII, fig. 17)

The specimens of this alga were very few in culture and collected only
once. They all measured the same, being 12 mu wide and 60.8 mu long, and
all were lunate in shape with the sharply pointed or spined tips bent in op-
posite directions. This is being tentatively identified as CI. lunula since the
description of that species fits with the exception of dimensions. CI. lunula
is 9-12 mu wide and 25-31 mu long.

Coll. June, 1937, drainage ditch in Haskell bottoms, Lawrence.

Thompson : Fresh-water Algae 35

Ankistrodesmus falcatus (Corda) Rolfs var. adcularis

(A. Braun) G. S. West

(PI. vir, fig. 19)

Cells solitary, 1.5-3.5 mu thick, 20-80 niu long, almost straight, but may

have a very slight arc. Chloroplast parietal and with or without a pyrenoid.

Coll. 1935, 1936, 1937, Lawrence. Comes up in any culture.

Ankistrodesmus falcatus (Corda) Rolfs var. mirabile W. and

G. S. West

(PI. VII. fig. 18)

Cells solitary, of the same measurements as the above variety, straight in
the main portion with the attenuated ends suddenly curved or they may be
spirally twisted, not quite making a complete turn.

Coll. 1935, 1936, 1937, LawTence. Comes up in any culture.

Schroederia setigera (Schroder) Lemm.

(PI. VII, fig. 13)

Cells solitary, with a long slender spine at each end, 2.5-10 mu wide, 22.5-
52.3 mu long without the spines, narrowly elliptic and sometimes slightly
lunate.

Coll. April, May, 1935, state lake, Tonganoxie.

Selenastrum gracile Reinsch.

(PI. VIII, fig. 1)

Cells arcuate to lunate with sharp poles, aggregated in groups of 4, 8, or 16
with their convex faces opposed. There is no colonial gelatinous sheath.
Cells 19-28 mu long, 4-5 mu wide.

Coll. June, 1937, Chetopa creek, Altoona.

Kirchneriella contorta (Schmidle) Bohlin

(PI. VIII, fig. 2)

Cells are irregularly curved cylinders, blunt on both ends, .7-2 mu wide, 8-10
mu long. Chloroplast single and parietal with one pyrenoid. Cells grouped
in fours or eights in a homogenous matrix.

Coll. May, 1935, April, 1936, state lake, Tonganoxie.

Kirchneriella lunaris (Kirchner) Mob.

(PI. VIII. fig. 3)

Cell.* lunate with the extremities pointed; 3-5 mu wide, 6-10 mu long.
Chloroplast single and parietal with one pyrenoid. Cells grouped in fours in
a colonial matrix.

Coll. May, 1936, Jackman's pond, Lawrence.

Tetraedron caudatum (Corda) Hansg.

(PI. VIII, fig. 4)

Cells solitary, five-lobed with a deep sinus, the lobes rounded and tipped
with a spine 3 mu long; 13-23 mu in diameter.
Coll. Sept., 1935, Jackman's pond, Lawrence.

36 The University Science Bulletin

Tetraedron punctulatum (Reinsch.) Hansg.

(PI. VIII, fig. 6)

Cells solitary, four-sided, the sides more or less equal, corners rounded
and the margin wavy. The corners may have the semblance of a short, blunt
spine. Cells 18-21 mu wide. Chloroplasts one to many, parietal and disk-
shaped, with or without a pyrenoid. Cell wall may be slightly warty.

Coll. May, 1936, Jackman's pond, Lawrence.

Tetraedron rmnimum (Al. Braun) Hansg.

(PI. VIII, fig. 5)
Cells more or less square, the wall smooth, the corners rounded and more
or less deeply indented between corners. Cells 6-10 mu wide, 3-6 mu thick.
Coll. March, 1937, Village creek swamp, Chanute.

Tetraedron trigonum (Naegeli) Hansg. Var. minor Reinsch

(PI. VIII. fig. 9)

Cells triangular, the sides indented and attenuating at the apices into sharp
spines, 10-14 mu wide.

Coll. July, 1937, drainage ditch in Haskell bottoms, Lawrence.

Tetraedron proteij orme (Turn.) Brunn.

(PI. VIII, fig. 11)

Cells two- or three-cornered, the corners drawn out into fine spines. Cells
twisted so that one pair of spines is opposite in direction to the other pair.
36-65 mu long.

Coll. July, 1937, stock pool in the sand hills, Burrton.

Tetraedron trilobatum (Reinsch) Hansg.

(PI. VIII, fig. 7)
Cells triangular, sides deeply indented and the corners broadly rounded.
Cells 25 mu wide.

Coll. July, 1937, stock pool in the sand hills, Burrton.

Tetraedron tumididum (Reinsch) Hansg.

(PI. VIII, fig. 10)
Cells tetrahedral, sides somewhat indented, corners broadly rounded. 20-60
nui wide.

Coll. June, 1937, Chetopa creek pool, Altoona.

Tetraedron pentaedricum W. and G. S. West

(PI. VIII, fig. 8)

Cells 5-Iobed, sides concave, lobes rounded and each bearing a short, stout,
slightly bent spine. Cells 10-15 mu in diameter without the spines. Spines
4.5-5.5 mu long.

Thompson : Fresh-watee Algae 37

Family Scenedesmaceae
Scenedesmus denticulatus Lagerh.

(PL VIII, fig. 18)

Cells egg-shaped to lance-elliptic with two or three teeth on each end.
Cells 4-11 mu wide, 6-15 mu long.

Coll. July, 1937, Chetopa creek pool, Altoona.

Scenedesmus obliqims (Tiirp.) Kiitz.

(PI. VIII, fig. 12)

Coenobium of four or eight cells in one or two rows. Cells pointed at
the ends, 4-35 mu long, 2.5-10 mu wide. Chloroplast parietal and contains
one pyrenoid.

Coll. 1935, 1936, 1937, in most any collection, Lawrence.

Scenedesmus bijuga (Turp.) Lagerh.

(PI. VIII, fig. 14)

Cells oval to ellipsoidal with broadly rounded ends; 4-8 in a coenobium in
one or two rows. Cells 4-7 mu wide, 7-18 mu long.
Coll. 1935, 1936, 1937, Jackman's pond, Lawrence.

Scenedesmus quadricauda (Turp.) Breb.

(PI. VIII, fig. 15)

Coenobium 4-celled, the cells ovoid with rounded poles. End-cells each
with two long spines. Cells 3-15 mu wide, 8-42 mu long.

Coll. Common in all cultures spring and summer, Lawrence.

Scenedesmus abundans (Kirch.) Chod.

(PI. VIII, fig. 16)

Coenobium 4-celled; cells ovoid and broadly rounded at the poles. End-
cells with 2-3 radiating spines between two long spines that are similar in
position to those of S. quadricauda. Cells 3-12 mu wide, 6-35 mu long.

Coll. April, 1935, Oct., 1936, Jackman's pond, Lawrence.

Scenedesmus dimorphus (Turp.) Kiitz.

(PI. VIII, fig. 13)
Cells long-pointed; the middle ones somewhat lens-shaped, the end ones
lunate; 6-7 mu wide, 30-40 mu long. Coenobium 4-celled.
Coll. May, 1935, 1936, Jackman's pond, Lawrence.

Crucigenia rectangularis (Nag.) Gay

^ (PI. VIII, fig. 17)

Cells in groups of four to form a coenobium which is free-floating. The
cells are quadrately arranged with a large space in the middle. They are
ellipsoidal in shape and contain from one to four parietal, discoid chloroplasts.
Cells 4-5 mu wide, 4-6 mu long.

Coll. Oct., 1936, Dightman's crossing, Wakarusa, Lawrence.

38 The University Science Bulletin

Crudgenia tetra'pedia (Kirch.) W. and G. S. West

(PI. VIII, fig. 23)

Coenobium 4-celled, generally joined into 16-celled compound coenobia.
Cells triangular and closely adhering to one another. The outer wall of the
cells is slightly concave. Cells 4.8 to 9.5 mu long.

Coll. July, 1937, Village creek, Chanute.

Tetrastrum elegans Playfair

(PI. VIII, fig. 22)

Coenobium 4-celled; the cells closely packed and each bearing a single long,
slender spine up to three times the cell in length. Cells 4.9 mu wide.
Coll. July, 1937, stock pool in sand hills, Burrton.

Tetrastrum apiculatum (Lemm.) Schmidle

(PI. VIII, fig. 21)

Coenobium 4-celled ; cells oblong, almost three-cornered, on the inside of the
outer poles bearing a veiy short, blunt spine. Cells 2.5-5 mu wide, 4-7 mu long.
Coll. July, 1937, stock pool in sand hills, Burrton.

Actinastrum gradllimum G. M. Smith

(PI. VIII, fig. 19)

Coenobium of 5, 8 or 16 cells; cells elongate and radiate in arrangement,
only slightly tapered at the poles, 3.3 mu wide, 9.9-13.2 mu long.
Coll. July, 1937, stock pool in sand hills, Bun-ton.

Actinastrum Hantzschii Lagerh. var. javanicum Bernard

(PI. VIII, fig. 20)
Coenobium of 4-16; cells elongate and radiate in arrangement, poles of the
cells tapering to a blunt or acute apex, 3.3 mu wide, 16.5 mu long.
Coll. July, 1937, stock pool in sand hills, Burrton.

Micractinium pusillum Fresenius

(PI. VIII, fig. 24)

Cells spherical, united into 4-celled coenobia which are always in turn united
with many coenobia to form multiple coenobia. The free face of each cell in
a coenobium bears from one to seven delicate setae which are many times
the cell diameter in length. Cells 6.6 mu in diameter.

Coll. July, 1937, stock pool in sand hills, Burrton.

Errerella bornhemiensis Conrad

(PL VIII, fig. 25)

Cells spherical, united into 4-celled coenobia in which the cells are pyram-
idal in arrangement. Multiple coenobia are the rule with over 100 cells, the
number usually being a multiple of four, 16, 64, to 256. The free face of
each cell bears a single, long, stout spine. Cells 3-7 mu in diameter.

Coll. July, 1937, drainage ditch in Haskell bottoms, Lawrence.

<

Thompson: Fresh-water Algae 39

Order SIPHONALES

Family Vaucheriaceae

Vaucheria geminata (Vaucher) D. C.

(PI. IX, fig. 1)

Thallus composed of irregularlj' branched, coenocytic filaments from which
the reproductive bodies are cut off by cross walls. Two oogonia are borne
laterally at the end of a short side branch with an antheridium rising between
them. Ripe oospores 64.5-190 mu long, 52.5-225 mu wide. Chloroplast dis-
coid, numerous and embedded in a parietal sheet of cytoplasm.

Coll. 1935, 1936. 1937, boat slough at north end of Ohio street, Lawrence.

Vaucheria sessilis (Vaucher) D. C.

(PI. IX, fig. 2)

Similar to the above species except that the antheridia and oogonia arise
next to each other and are sessile on the main filament. The particular material
found is Pascher's jorma clavala, which has erect oogonia with the receptive
spot on the top. Oospores 66-88.5 mu long, 49.5-66.5 mu wide.

Coll. April and May, 1937, Jackman's pond, Lawrence.

Order ZYGNEMATALES

Family Zygnemataceae

Zygnema in^igne (Hassall) Kiitz.

(PI. IX, fig. 3)

Vegetative cells 26-30 mu wide. Conjugation either lateral or scalariform,
the cells concerned swollen or only the cell acting as zygote container swollen.
Zygospore ovoid, 26 mu wide, 32 mu long. Exospore thin, smooth and color-
less. Mesospore thick, brown and smooth.

Coll. Sept., May, 1935, 1936, quarry pool on East Fifteenth street, Lawrence.

Spirogyra protecta Wood

(PI. IX, fig. 4)

Vegetative cells 20-25 or 36-40 mu wide. Conjugation scalariform. One
chloroplast is present in each cell. Cross walls replicate. Neither of the
conjugating cells becomes swollen. Zygote ovoid; exospore thick, colorless
and of two layers. The outer layer thin and smooth, the inner thick and
ornamented with circular depressions which make the intervening areas stand
out as spines. Zygote 36 mu wide, 72 mu long.

Coll. April, 1936, cjuarry pool on East Fifteenth street, Lawrence.

Spirogyra punctiformis Transeau

(PI. IX, fig. 6)

Vegetative cells 27-30 mu wide, with one or two chloroplasts, end walls
not replicated. Cell containing the zygote becomes very swollen. Zygote 40-48
mu wide, 60-110 mu long; exospore colorless, thick and smooth, mesospores
fairly thick, yellow and minutely pitted.

Coll. April, 1935, Lawrence.

40 The University Science Bulletin

Spirogyra malmeana Hirn

(PL IX, fig. 5)

Vegetative cells 76-91 mu wide, end-walls not replicated, three or four
chloroplasts to a cell. Conjugation scalariform, the cells containing the zygote
not becoming swollen. Zygote ellipsoidal, 54 mu wide, 82 mu long. Exospore
thin, smooth and colorless. Mesospore, thick, brown and ornamented with
irregular and branched ridges which do not form a net on the outer surface
of the mesospore. There are peculiar crystals present in the cytoplasmic
strands that are shaped like a cross.

Coll. April, May, 1936, Potter's pond, Lawrence, roadside ditch, Lecompton.

&jnrogyra decimina (Miill.) Cziirda emend.

(PI. IX, fig. 13)
Vegetative cells 31-33 mu wide, with replicate end-walls, and with one
chloroplast. Conjugation scalariform, the canal being formed largely by one
of the conjugating cells. Zygote containing cell scarcely swollen. Zygote 34-62
mu wide, 56 to 124 mu long. Exospore thin, smooth and colorless. Mesopores
thick, yellow and marked with an equatorial line running lengthwise.

Coll. June, 1937, boat slough at north end of Ohio street, Green's pond,
Lawrence.

Family Mesotaeniaceae

Spirotaenia parvula Arch.

(PI. IX, fig. 7)

Cells solitary, fusiform, with a single, parietal and spirally twisted chloro-
plast. Cells 25 mu wide, 145 mu long.

Coll. May, 1936, East Fifteenth street, quarry pool, Lawrence.

Netrium digitus (Ehrenb.) Itz. and Rotlie

(PI. IX, fig. 9)

Cells large, 216-264 mu long, 69-72 mu wide, oblong-elliptic, gradually at-
tenuated from the middle to the rounded or rounded-trimcate apices. Chloro-
plast axial with 5-7 radiating plates which are deeply serrate on the outer
margins, two to a cell. Cell wall smooth.

Coll. June, 1937, Moulton's lake, Neodesha.

Roya obtusa (Breb) W. and G. S. West

(PI. IX, fig. 8)

Cells cylindrical, very slightly curved, apices broadly rounded, 22-120 mu
long, 12-15 mu wide. Chloroplast a long band the entire length of the cell
and containing 2-8 pjTenoids, and with a notch at the middle.

Coll. June, 1937, Chetopa creek, Altoona.

Family Desmidiaceae
Closterium moniliforme (Borg.) Ehrenb.

(PI. IX, fig. 12)
Cells solitary, lunate, poles attenuated to blunt points. Chloroplasts with
longitudinal ridges and a row of pyrenoids. Cells 291 mu from tip to tip and
55.7 mu wide at the isthmus.

Thompson: Fresh-water Algae 41

Coll. 1935, 1936, 1937, spring, abundant in Green's pond, Jackman's pond,
East Fifteenth street quarry pool, Lawrence.

Closterium acerosum (Schrank) Ehrenb.

(PI. LS, fig. 11)

Cells linear-lanceolate with the ends very slightly curved; 343 mu long,
25.7 mu wide at the isthmus.

Coll. May, 1936, Green's pond, Lawrence.

Closterium subulahim (Kiitz.) Breb.

(PI. IX, fig. 10)
Cells small, slightly curved, 148-150 mu long, 9-11 mu wide; apices gradually
attenuated from the middle, subtruncate to rounded on the end. Cell wall
smooth. Chloroplast with three pyrenoids.

Penium curciibitum Biss.

(PI. IX, fig. 15)

Cells aggregated in a copious jelly. Individual cells with a very shallow
constriction at the isthmus, wall smooth ; 38 mu long, 19 mu wide. Chloroplast
in each semicell two or four lobed. Cells ovate-elliptic.

Coll. April, 1937, Santa Fe pond, Chanute.

Pleurotaenium Ehrenbergii (Breb.) De Bary

(PI. IX, fig. 14)

Cells linear with truncated apices and with a constriction at the isthmus,
384-437 mu long, 30-33 mu wide. Semicells inflated and undulated at the
base. Cell wall punctate, apices with 7-8 tubercles in a ring about the margin.
Chloroplasts numerous, parietal, longitudinal bands, each containing many
pyrenoids.

Coll. June, 1937, Chetopa creek, Altoona.

Euastrum verrucosum Ehrenb.

(PL IX, fig. 22)

Cells moderately large, subhexagonal, somewhat longer than broad, deeply
constricted, sinus narrowly linear; semicells three-lobed, interlobular incisions
deep but open; polar lobe widely cuneate, angles rounded and granulate,
apex retuse; lateral lobes about as wide as the polar lobe, cuneate and bilobu-
late, lower lateral lobule subconical, rovmded, granulate, and horizontally
directed, upper lateral lobule granulate, divergent upward and outward; semi-
cells with three large protuberances across the broadest part, the central one
the largest, each protuberance with large granule.s in concentric circles; cell
wall granulate. Cells 84-90 mu long, 72-81 mu wide.

Coll. June, 1937, Chetopa creek, Altoona.

Cosmarium circulare Reinsch

(PI. IX, fig. 19)
Cells orbicular with two narrow sinuses. Apex of the sinus not inflated;
cell wall not ornamented. Cells 48.3 mu long and nearly the same in width.
Coll. Oct., 1935, East Fifteenth street quarry pool. Lawrence.

42 The University Science Bulletin

Cosmarium punctulatum (Nordst.) Borg.

(PI. IX, fig. 18)
Cells orbicular, flattened at the poles; minutely scalloped in outline, punc-
tate with groups of punctae on the cheeks. Cells 33.3 mu long, 30 mu wide.
Coll. May, 1936, boat slough at north end of Ohio street, Lawrence.

Cosmarium renijorme (Ralfs) Arch.

(PI. IX, fig. 16)

Cells flattened-orbicular, 45.6-55 mu long, 49.4-54 mu wide. Semicells reni-
form in shape so that a definite pore is seen, on each side of the isthmus,
which is triangular in shape. Wall covered with coarse granules in rows. In
side view each semicell is circular, in vertical view elliptic with slight swellings
on each side at the middle.

Coll. April, 1936, Green's pond, Lawrence; June, 1937, Moulton's lake,
Neodesha.

Cosmarium polymorphum

(PI. IX, fig. 17)

Cells orbicular with the poles flattened and the wall at these places smooth
and even. The remainder of the wall on each semicell from the pole to the
sinus is crenate in outline. Sinus inflated slightly at the apex. Cells 8 mu
long, 8 mu wide.

Coll. May, 1936, Lawrence.

Cosmarium Schliephackeanum Griin. (Taft 1931)

(PI. IX, fig. 21)

Cells small, angular, 15 mu long, 12-15 mu wide, deeply constricted, sinus
narrowly linear, not inflated at the apex; semicells transverselj^-hexagonal,
angles not rounded, apex broadly truncate. Vertical view narrowly elliptic
with dull or sharp protuberance on each side at the middle. One chloroplast
and one p>Tenoid in each semicell.

Coll. May, June, 1936, Jackman's pond, Lawrence.

Cosmarium pseudobroomei Wolle (Hylander 1928)

(PI. IX, fig. 20)

Cells square to rectangular with the angles rounded, 33 mu long, 38-40 mu
wide, 13 mu thick, walls densely granular with very small granules, deeply
constricted, sinuses narrow and dilated at the apices. Semicells rectangular
in outlines, angles rounded, sides straight or slightly convex, apex straight.
Lateral view; semicells orbicular. Vertical view; rectangular-elliptic. Chloro-
plast in each semicell, axial and containing two pyrenoids.

Coll. June, 1937, Green's pond, Lawrence; July, 1937, Moulton's lake,
Neode.sha.

Cosmarium polygonum (Nag.) Arch.

(PI. X, fig. la)

Cells 21 mu long, 15 mu wide, 5.7 mu thick, deeply constricted, sinus narrow
and not dilated at the apex. Semicells six-sided in outline, angles rounded,
sides and apex slightly concave. One axial chloroplast and one pyrenoid in
each semicell. Lateral view narrowly or linear elliptic with the poles rounded-
truncate. There is a sharp inflation bearing a granule, on each side a little

Thompson: Fresh-water Algae 43

above the constriction. Vertical view narrowly elliptic with the poles either
rounded or rounded-truncate, a sharp inflation and granule in the middle on
each side.

Coll. July, 1937, Chetopa creek, Altoona.

Two other specimens of Cosmarium were collected which, though not
fulfilling the dimensional requirements of the description of C. Polygonum
and not the same shape, seem to be variants of C. polygonum. They each
have the same type of inflation or granular protuberance, sinuses, chloro-
plast and cell wall as C. polygonum and differ only in shape.

Variant 1
(PI. X, fig. lb)
Cells rectangular to square with the corners cut off, 19 mu long, 15 mu wide,
5.7 mu thick. Semicells six-sided in outline, the corners rounded, wall smooth.
One chloroplast and one pyrenoid in a semicell. Sinuses deep, narrow and
not inflated. Lateral view : semicells rectangular with the corners rounded
and the broad faces just above the constriction inflated and bearing a blunt
protuberance. Vertical view: rectangular with the corners rounded; middle
inflated and bearing a blunt protuberance in the center on each side.
Coll. June, 1937, Altoona.

Variant 2

(PI. X, fig. ic)

Cells rectangular and sharp-angled, 11.4 mu long, 9.5 mu wide. Semicells
hexagonal with sharp corners, containing each one chloroplast and one pyre-
noid. Sinuses deep and not inflated at the apices. Lateral view: semicells
ovoid, each with a centrally located blunt point on the broad face. Vertical
view: elliptic with the poles rounded-truncate and a blunt point in the center
on each side.

Coll. June, 1937, Neodesha.

Cosmarium granatum Breb.

(PL X, fig. 2a)

Cells subrhomboid-elliptic, 27-45 mu long, 21-31 mu wide, 12 mu thick,
wall smooth. Semicells subpyramidate, corners rounded, poles rounded-trun-
cate or rounded, one chloroplast and one pyrenoid in each. Sinuses deep,
narrow and slightly dilated at the apex. Lateral view: semicells ovate-
pyriform, the whole narrowly elliptic. Vertical view: elliptic to slightly rhom-
boidal-elliptic. Cell wall very finely punctate.

Coll. July, 1937, Village creek, Chanute.

A number of specimens of Cosmarium were collected which seem to be
variants of C granatum. They differ from each other and from C. granatum,
in size and shape only. Descriptions of them are given below and they are
figured on the plate along with C. granatum.

Variant 1

(PI. X, fig. 2b)

Cells rhomboidal in outline, 27.5 mu long, 20 mu wide, 11.4 mu thick, wall

smooth. Semicells angularly heart-shaped, six-sided in outline, the corners

rounded, the poles rounded-truncate. Each semicell contains one chloroplast

•14 The University Science Bulletin

and one pyrenoid. Sinuses deep and slightly dilated at the apices. Lateral
view: semicells heart-shaped to ovate-pyriform. Vertical view: rhomboidal-
elliptic.

Variant 2

(PI. X, fig. 2c)

Cells ovoid, 19 mu long, 15 mu wide, 7.6 mu thick, wall veiy finely punctate,
deeply constricted, sinuses narrow and slightly dilated at the apices. One axial
chloroplast and one pyrenoid in each semicell. Semicells somewhat reniform
with the lateral walls straight and forming an angle with the sinuses and the
apical part of the semicell which is broadly rounded. Lateral view : semicells
ovoid. Vertical view: elliptical.

Coll. June, 1937, Moulton's lake, Neodesha.

Cosmarium sexpapillosum sp. nov.

(PI. X, fig. 4)

Cellula est rhomboidalis ovata; 77.9 mu longa, 57 mu lata; semicellulae
paulum triangulares, polls truncatis et paribus, lateribus leviter undulatis et
in polum vergentibus. Malae sunt ornatae sex magnis papillis, trinis in duobus
ordinibus dispositis. Binae chlorophorae et pyrenoides sunt singulis semi-
cellulis; sinus alti sunt et in apicibus inflati.

Cells rhomboid-ovate, 77.9 mu long, 57 mu wide, 28 mu thick, 14.5 mu wide
in the isthmus. Semicells somewhat angularly three-sided, the poles truncate
and even, the margins slightly crenate, cheeks ornamented with six prominent
granules that are arranged in two rows of three each. Chloroplasts and py-
renoids two to each semicell. Sinuses deep and inflated at the apices.

Coll. spring, 1935, 1936, 1937, boat slough at the north end of Ohio street,
Lawrence,

Cosmarium impressidum Elfv. (Taylor 1934)

(PL X, fig. 3)

Cells somewhat rectangular with the poles slightly inflated, 21-26 mu long,
14-18 mu wide, isthmus 4 mu wide. Semicells obscvn-ely six-sided, margins
undulate, lateral walls slightly divergent. The material collected measured
19 mu long, 15.2 mu wide, 9.5 mu thick, isthmus 5 mu wide. Sinuses deep
and slightly inflated at the apices. One chloroplast and one pyrenoid in each
semicell. There is a conspicuous, blunt granule in the center of each face
of the semicells. Top view elliptic with a median granule on each side.
I^ateral view: semicells nearly orbicular to ovoid with the granule on each
side median in position.

Coll. June, 1937, Moulton's lake, Neodesha.

Cosmarium Nymannianum Gvxm.l (Taylor 1934)

(PI. X, fig. 5)

The Cosmarium that is being tentatively determined as C. nymannianum
is possibly a variety. Its measurements are 37.9-39.6 mu long, 29.7 mu wide,
13.2-19.8 mu thick, and the isthmus is 10 mu wide. Semicells are broadly
heart-shaped. Sinuses are deep and very slightly inflated at the apices. There
is a small pore in the center of each face of a semicell which is more evident
in the lateral and top views. This pore is difficult to see except in cells in

Thompson: Fresh-water Algae 45

which the contents have disintegrated. In lateral view the semicells are
narrowly heart-shaped to ovate. The wall is smooth, but thickened about
the pore. In vertical view the semicells are elliptic with the pore evident
in the thickened portion of the cell wall median in position on each side.
The cells of this material are approximately half the size of C. nymannianum.
the dimensions of which are : 52-54 mu long, 38-42 mu wide, 24-26 mu thick
and 7.5-9.5 mu wide at the isthmus. There is some difference in the shape
of the semicells; those of the present material have the lateral walls regularly
convergent towards the pole, those of C. nymanniamum have the lateral walls
slightly depressed. Both have the pore in the face of each semicell. This
material falls well within the dimensions of Cosmarium granatum, but the
latter species has no pores.

Coll. June, 1937, Green's pond, Lawrence.

Cosmanum humile (Gay) Nordst. var. lacustre Taylor, Wm. R.

(Taylor 1934)

(PI. X, fig. 6)

Cells very small, about as long as broad; semicells subrectangular to trapezi-
form, the basal and upper angles obliquely truncate, the sides bidentate,
the upper indentation deeper; apex about six-crenate with a slight projection
of the ridges, especially the submarginal ones, on to the lateral faces; isthmus
linear; face of semicell usually with two low swellings above the basal angles,
or these obsolete, and with strong costae, the lateral ones somewhat weaker
than the central; vertical view inflated-oval, the top faintly ridged, the sides
crenate because of the costae; walls thin except on the incrassate faces.
Length 14-15 mu, width 15-16 mu, thickness 8.5 mu, isthmus 5 mu.

The specimens collected coiTespond to Taylor's description with the ex-
ceptions that the margins are more crenate than dentate, there are only four
small crenations on the apex of a semicell, the costae are more distinct, be-
ing composed of a compoimd of three ridges, the central one of which is
square when seen from above and the two outer are rounded; and, that the
dimensions of the collected material are slightly greater. Length 22.8 mu,
width 19 mu, thickness 11.4 mu, isthmus 7 mu.

Coll. July, 1937, Moulton's lake, Neodesha.

Cosmarium crenatum Rolfs (Hy lander 1928)

(PI. X, fig. 7)

Cells rectangular with the angles broadly rounded, 36.6 mu long, 23.9 mu
wide, 14.8 mu thick, deeply constricted, sinuses narrow with apices quite
dilated. Semicells subquadrate, the lateral walls parallel half way then gradu-
ally rounding into the flat pole. Polar wall very minutely crenated, lateral
walls broadly crenated to undulate; faces ornamented with many small un-
dulate inflations arranged in rows radiating from the poles. Chloroplast single,
reticulate and parietal in each semicell. Lateral view: rectangular with the
corners rounded, semicells rhomboidal. Vertical view: rectangular-elliptic.

Coll. June, 1937, Moulton's lake, Neodesha.

4G The University Science Bulletin

Micrasterias americana (Ehrenb.) Ralfs.

(PI. X, fig. 8)

Cells hexagonal in outline, 132-150 mu long, 112-120 mu wide, deeply con-
stricted, sinuses acute at the apices, opening outwards, the margins sometimes
undulate and closing. Semicells five-lobed, end lobe about as wide as the
combined lateral pair, expanded from the base outward, apical margins
widely concave, each of the four angles formed ending in a thick divergent
process with their apices truncately toothed; incisions on each side of the
polar lobe are deep and open; two small papillate projections occur, one on
each side of the base of the incision; lateral lobes separated by a shallow
incision whose apex is rounded; each lobe divided again into two smaller
lobes by an open shallow incision, each small lobe with four to seven acute
teeth; chloroplast axial, five-lobed, end lobe concave or incised.

Coll. June, 1937, Chetopa creek, Altoona.

Staurastrum paradoxmn Meyen

(PI. X, fig. 9)

Semicells with three radiating arms that are annulately ringed with minute
spines and which end truncately with divergent spines. Semicells contain a
single chloroplast which radiates into the arms; it is axial and contains one
pyrenoid.

Coll. April, 1937, Green's pond, Jackman's pond, Lawrence.

CLASS DINOPHYCEAE
SUBCLASS DINOFLAGELLATAE

Glenodinium oculatiim Stein

(PI. X, fig. 10)

Equatorial furrow completely around the cell, sulcus obscure. Chloroplasts
few to many, discoid, greenish-.vellow in color. The halves of the cell are
approximately equal. Length 20-23 mu, width not given. The material col-
lected measured 19-20 mu long and 15.2 mu wide.

Coll. March, 1937, boat slough at north end of Ohio street, Lawrence.

Glenodinium pulvisculus Stein

(P!. X, fig. 11)
Cells 23 mu long, 18.4 mu wide. Cells rhomboidal in outline. Cell halves
broadly bell-shaped. Walls of the equatorial furrow overhanging. Sulcus
a very slight longitudinal furrow. The material collected measured 17-20 mu
long and 16 mu wide.

CLASS EUGLENOPHYCEAE

Family Euglenaceae

Euglena intermedia (Klebs) Schmitz

(PI. X, fig. 12)
Cells linear-cylindric with a short suddenly-pointed posterior end, 120-135
mu long, 8-12.5 mu wide. Membrane delicately, spirally striated. Flagellum
short. Chloroplasts numerous, discoid, without pyrenoids. Paramylum bodies

Thompson: Fresh-water Algae 47

numerous throughout the ceil, rectangular with smoothly rounded ends. Di-
viding cells secrete a gelatinous sheath about them.

Coll. 1935, 1936, 1937, year around, drainage ditch in Haskell bottoms,
Green's pond, boat slough at north end of Ohio street, Lawrence.

Euglena spirogyra Ehrenb.

(PI. X, fig. 13)

Cells weakly metabolic, lanceolate-cylindric, the forward end less attenuated
than the rear, the rear end veiy gradually attenuated into a fine hyaline,
rigid point, 80 mu long, 8 mu wide. Membrane yellow to dark brown, with
spirally arranged beaded ridges, spiralling to the left. Flagellum very short.
Chloroplasts numerous, discoid, dark green and without pyrenoids. Two
large, thick paramylum rings lie, one before and one after the nucleus. Neu-
cleus central in position.

Coll. March, April, 1936, June, July, 1937, drainage ditch in Haskell bottoms,
Lawrence.

Euglena acus Ehrenb.

(PI. X, fig. 14)

Cells rigid or very slightly metabolic, spindle-shaped, linear with a truncated
anterior pole, the posterior end gradually attenuated into a long slender point,
140-180 mu long, 10 mu wide. Chloroplast numerous, discoid and lack py-
renoids. One to several paramylum bodies present.

Coll. 1935, 1936, 1937, April to July, Green's pond, Lawrence.

Euglena rubra Hardy

(PI. X, fig. 15)

Cells ob-lanceolate to cylindric with a short blunt end-point at the pos-
terior end, up to 200 mu long, 38-60 mu wide, colored a deep red-brown by
haematochrome. Membrane spirally striated to the left. Flagellum about
three quarters the length of the cell. Chloroplasts numerous and discoid.
Paramylum grains numerous, short cylindric. Dividing cells remain for a
time within the greatly enlarged cyst, aggregates of dividing cells form a scum
on the water surface pseudoparenchymatous in appearance.

Coll. June, July, 1937, East Fifteenth street quarry pool, Lawrence.

Euglena truncata Walton* var. baculifera var. nov.

(PI. X, fig. 16)

Cellula est cylindrata, 188-198 mu longa, 19.8-23 lata; anteriore polo acutiore,
posteriore truncato. Membrana est ad sinistram forma spirae striata et a
truncato polo spissata. Flagellum par est tertiae parti corporis. Nucleus
ovatus in medio corporis positus est. Paramylum est simile longo et tenui
baculo a tertia parte flexo. Pyrenoides desunt. Chlorophorae sunt multae
et discoides.

Motile cells cylindric, 188-198 mu long, 19.8-23 mu wide, anterior end
fairly pointed, posterior end truncated and .slightly concave. Periplast spirally
striated to the left, thickened in the truncated posterior portion. Eyespot
just posterior to the conspicuous gullet ; fiagellum approximately one third

Walton, 1915.

48 The University Science Bulletin

the length of the body. Nucleus oval and variable in position, being slightly
anterior, central or slightly posterior. Chloroplasts numerous, discoid, slightly
longer than broad. No pjTenoids. Paramylum in the form of a single
long stick which may be slightly bowed but more characteristically is bent
at one third of its length at about an angle of 165 degrees. It lies anterior
to the nucleus when the nucleus is posterior in position; posterior to the
nucleus when the nucleus is anterior in position and either above or below
when the nucleus is central in position. The paramylum stick is 4-6.6 mu
thick and 52.7-66 mu long. There is a large vacuole in the posterior end which
becomes more evident when the organism is exhibiting euglenoid movement.
"V\Tiile the characteristic swimming form is cylindric, the organism varies a
bit and while swimming may become broadened and flattened at the pos-
terior end or both anterior and posterior ends. When both ends are flattened
it generally becomes slightly twisted to the left. The partial rigidity of the
periplast of the truncated portion causes the formation of a depression at
the posterior pole. In resting condition the organism becomes very broad and
flattened, elliptic in outline. Those measured in this condition varied from
115.5-151.8 mu long and 37.9-42 mu in width. Encj-stment and division were
not observed.

E. tnmcata var. bacuUfera is quite similar to E. truncata Walton (Walton,
1915) in general cell shape and in the truncated posterior end. The lengths of
the two organisms are close enough to come under variability, E. truncata be-
ing 175 mu long, 27-29 mu wide, and var. bacuUfera being 188-198 mu long and
19.8 to 23 mu wide. However, Walton describes E. truncata as having the nu-
cleus posterior in position and as having numerous small paramylum grains and
no large ones. Walton also makes the statement that E. truncata appeared in
almost all cultures taken over a period of three years, so his organism must
have been fairly constant in the characters he has described for it. The pres-
ent variety, though not collected over as long a period, was constant in its
characters for all the collections taken in one summer. There were several
accessions during the summer on account of rains, so there was good chance
for any variability to appear during that season. Since the lack of small
paramylum bodies and the presence of one large characteristic body has been
constant for the present organism and since the truncated posterior end is so
remarkably different from all other species of Euglena the author feels that
the present organism should be described as a variety of E. truncata rather
than as a distinct and new species.

Coll. summer, 1937, Village creek swamp, Chanute.

Euglena alata sp. nov.

(PI. X, fig. 17)

Cellula est similis fasciae, 165-209 mu longa, 22.8-23 mu lata; anteriore parte
rotunda, posteriore in brevem pinguem caudam spinae similem subito attenu-
ata; toto corpore ad dextram torto. Membrana est in longitudinem striata,
et elata in formam trium pinnarum a medio ad posteriorem parlem perti-
nentium. Flagellum circiter par est tertiae parti corporis. Nucleus ovatus in
medio corporis positus est. Paramylum consistit ex duobus amplis anulis, uno
ante nucleum, altero post nucleum. Pyrenoides desunt. Chlorophorae sunt
multae et discoides.

Thompson : Fresh-water Algae 49

Motile individuals 165-209 mu long, 22.8-23 mu wide, verj^ slightly spirally
striated to almost longitudinally striated and characteristically twisted to the
right. The anterior portion is flattened to form two broad faces. Along the
edge of the flattened portion and opposite to the eyespot there is a groove
which broadens out posteriorly so that from the middle to the posterior end
there are formed three flanges with concave faces in between. The eyespot
lies some distance back from the anterior end and next to the posterior end
of the gullet. The flagellum is shorter than the body, approximately one third
the body length. The nucleus is oval and central in position. Two long, nar-
row paramylum rings are present, one anterior to and the other posterior to
the nucleus. No pyrenoids are present. Chloroplasts, small, numerous and
discoid. The posterior end attenuates abruptly into a stout spinelike tail.

It is felt that those individuals measuring 165-189 mu in length are juvenile
forms and that 209 mu is the upper limit of length, since a number of dividing
individuals as well as numerous motile ones were of this length. Prior to di-
vision a motile individual comes to rest, becomes cylindrical and vacuolate and
finally divides longitudinally. No gelatinous sheath or division cyst is formed.

The two described species nearest to Eugletm alata are E. oxyuris and E.
tripteris. E. oxyuris is 375-490 mu long, 30-45 mu wide and is figured as
cylindrical in shape. E. tripteris is 70-80 mu long, 8-14 mu wide and is figured
as twisted and flanged in shape. Euglena alata falls well above the greatest
length of E. tripteris and well below the shortest length of E. oxyuris.

Coll. June, July, 1937, drainage ditch in Haskell bottoms, Lawrence; Village
creek swamp, Chanute; Chetopa creek, Altoona.

Euglena fusca (Klebs) Lemm

(PI. X, fig. 21)

Cells weakly metabolic, elongate and flattened, posterior gradually attenu-
ated and ending in a short stout tail, 90 to 225 mu long, 23-27.5 mu wide.
Periplast dark brown to black, longitudinally striated. Flagellum as long as
the body. Chloroplasts discoid, numerous. Pyrenoids lacking. Two large,
thick paramylum rings are present, one on each side of the nucleus. Divid-
ing and resting cells without a gelatinous sheath.

Coll. June, July, 1937, drainage ditch in Haskell bottoms. East Fifteenth
street ditch, Lawrence.

Euglena tripteris (Duj.) Klebs

(PI. X, fig. 23)

Cells slightly metabolic, long band-shaped, spirally twisted, with a long,
slender, sharp tail, 70-80 mu long, 8-14 mu wide. Periplast longitudinally
striated. Flagellum one half the body length. Chloroplasts numerous, dis-
coid. Pyrenoids lacking. There are two long sticklike paramylum bodies, one
before the one back of the nucleus. Division stage sheathless.

Coll. June, July, 1937, Green's pond, Lawrence; June, Chetopa creek pool,
Altoona.

Euglena torta Stokes

(PI. X, fig. 24)

Cells rigid, spindle-shaped, spirally twisted to the left, with a sharp tail, 63
mu long, 13 mu wide. Membrane smooth, flagellum almost body length.

4—4141

50 The University Science Bulletin

Chloroplasts numerous, discoid. Pyrenoid? There are two long sticklike para-
mylum bodies, one before and one behind the nucleus. Division and resting
stages not known.

Coll. July, 1937, Green's pond, Lawrence.

Euglena acutissima Lemm.

(PI. X, fig. 28)

Cells rigid, linear-spindle-shaped, attenuated into a long sharp tail, 123 mu
long, 7 mu wide. Periplast weakly, spirally striated. Flagellum short. Chloro-
plasts numerous, discoid, in spiral lines. Pyrenoids lacking. Two paramjdum
bodies are present, one before the other, behind the nucleus. Division and
resting stages not known.

Coll. May, June, 1937, Green's pond, boat slough at north end of Ohio
street, Lawrence.

Euglena deses Ehrenb.

(PI. X, fig. 18)

Cells very metabolic, long-cylindric or ribbonlike, with a veiy short tail,
80-90 mu long, 15-22 mu wide. Periplast w^eakly spirally striated to the left.
Flagellum short, chloroplasts numerous, discoid. Pyrenoids naked. Paramy-
lum bodies rodlike. Dividing cells within a gelatinous sheath.

Coll. Jul}', 1937, drainage ditch in Haskell bottoms, Lawrence.

Euglena fiava Dang.

(PL X, fig. 29)

Cells metabolic, spindle-shaped, with a short tail, 60 mu long, 25-30 mu
wide, colored red by the presence of haematochrome. Periplast smooth.
Flagellum body length. Chloroplasts 3-15, discoid. Pyrenoids doubly sheathed.
Paramylum? Dividing cell within a thin sheath.

Coll. July, 1937, Green's pond, drainage ditch in Haskell bottoms, Lawrence.

Euglena variabilis Klebs.

(PI. X, fig. 27)

Cells very livelj'-metabolic, short-cylindric with a short, blunt tail or al-
most egg-shaped. Periplast strongly, spirally striated, 30.5-46 mu long, 9-13
mu wide. Flagellum two to three times the body length. Chloroplasts nu-
merous, discoid. Pyrenoids lacking. Eyespot very large, dark red. There is
one large paramylum body near the gullet. Dividing cells egg-shaped, with-
out a sheath.

Coll. July, 1937, boat slough at north end of Ohio street, drainage ditch in
Haskell bottoms, Lawrence.

Euglena geniculata Duj.

(PI. X, fig. 22)

Cells livelj'-metabolic, long-cylindric, posterior portion attenuated into a
short tail, 70-85 mu long, 12-22 mu wide. Periplast delicately spirally striated.
Flagellum almost body length. Chloroplasts star-shaped, one before and
one behind the nucleus. Pyrenoids covered with a sheath of small paramylum
bodies. Dividing cells within a thin sheath.

Coll. July, 1937, Chotopa creek pool, Altoona.

Thompson : Fresh-water Algae 51

Euglena spiroides Lemm.

(PI. X, fig. 19)

Cells little metabolic, elongate-cylindric to ribbon-shaped, spirally twisted,
with a short tail, 60-170 mu long, 16 mu wide. Periplast delicatelj^ longitudi-
nally striated. Flagellum short. Chloroplasts numerous, discoid. Pyrenoids
lacking. Paramylum bodies small, round. Division and resting stages not
known.

Coll. May, 1936; Julj^, 1937, Green's pond, boat slough at north end of
Ohio street, Lawrence.

Euglena Grisoli Deflandre
(Pascher 1929)

(PI. X, fig. 20)

Cells weakly metabolic, long-cylindric, suddenly constricted at the posterior
end into a short tail, 68-75 mu long, 10-13 mu wide, spirally twisted or curved
to the left. Periplast spirally striated to the left with embossed, elongated,
bacteria-like verrucae. Chloroplasts numerous discoid. Two large sticklike
paramylum bodies are present, one above and one below the nucleus. Py-
renoids lacking. Flagellum approximately half the body length.

Coll. Jime, July. 1937, Green's pond, Lawrence.

Euglena viridis Ehrenb.

(PI. X, fig. 25)

Cells lively-metabolic, spindle-shaped with a short tail, 52-57 mu long, 14-18
mu wide. Periplast delicately spirally striated. Flagellum body length.
Chloroplast star-shaped, in front of the nucleus. Pyrenoids sheathed with
small paramylum grains. Dividing cells within a gelatinous sheath. Resting
cells within a thick, layered cyst membrane.

Coll. July, 1937, Green's pond, Lawrence.

Euglena proxima Dang.

(PI. X, fig. 26)
Cells lively-metabolic, spindle-shaped with a colorless tail, 60-70 mu long,
20 mu wide. Periplast spirally striated to the left. Flagellum one to one
and one half times the body length. Chloroplasts numerous, discoid. Py-
renoids lacking. Paramylum bodies ring-shaped or short-cylindric. Dividing
cells with a thin sheath. Resting cells with a thicker, layered membrane.
Coll. June, July, 1937, Green's pond, drainage ditch in Haskell bottoms
Lawrence.

Lepocinclis Steinii Lemm.

^ (PI. XI, fig. 6)

Cells short-spindle-shaped, 22-30 mu long, 8-15 mu wide. Posterior attenu-
ated into a tail 1.5-4 mu long. Periplast striated scarcely spirally almost
longitudinally. Two large paramylum rings present, one on each side of the
middle.

Coll. Ma3', June, July, 1937. Green's pond, boat slough at north end of
Ohio street, Lawrence.

52 The University Science Bulletin

Lepocinclis Butschlii Lemm. var. angustata Deflandre

(Pascher 1929)

(PI. XI, fig. 4)

Cells broad and plump, spindle-shaped to rhomboidal, 32-42 mu long, 13-16
mil wide. Periplast spirally striated to the left.

Coll. June, July, 1937. boat slough at north end of Ohio street, Lawrence.

Lepocinclis ovum (Ehrenb.) Lemm.

(PI. XI, fig. 1)

Cells oval, 30-38 mu long, 15-18 mu wide. Posterior end sharply attenu-
ated into a tail 6-7 mu long. Flagellum twice the body length. Periplast
strongly spirally striated to the left.

Coll. spring, July, 1937, boat slough at north end of Ohio street, Lawrence;
June, Chetopa creek pool, Altoona.

Lepocinclis ovum (Ehrenb.) Lemm. var. globula (Perty) Lemm.

(PL XI, fig. 2)

Cells almost orbicular, 20-27 mu long, 16-21 mu wide. Flagellum 2-3 times
the body length.

Coll. July. 1937, Green's pond, Lawrence.

Lepocinclis ovum (Ehrenb.) Lemim. var. palatina Lemm.

(PI. XI. fig. 3)

Cells egg-shaped, 20 mu long, 15.5 mu wide. Periplast spirally striated to
the left with short striae.

Coll. July, 1937, stock pool in the sand hills, Burrton.

Lepocinclis acicularis France

(PI. XI, fig. 5)

Cells long and small spindle-shaped, 21-22 mu long, 6-9 mu wide. Flagellum
over body length. Periplast spirally striate to the left with, at the most, 12
striae.

Coll. July, 1937, field ditch, Neodesha.

Lepocinclis fusiformis (Carter) Lemm.

(PI. XI, fig. 8)

Cells short and broad spindle-shaped or lemon-shaped, 25 to 51 mu long,
12-49 mu wide. Periplast delicately spirally striated to the left. Flagellum
1-1% times the body length. Anterior pole is a short snout which is divided
into two lips or papillae between which the flagellum is inserted.

Coll. July, 1937, boat slough at north end of Ohio street, Lawrence.

Lepocinclis turbinifonnis Deflandre
(Conrad 1934)

(PI. XI, fig. 7)
Cells turbinate, the anterior portion nearly spherical, attenuated rapidly
to a base fiom which there projects a short, pointed tail. Periplast spirally
striated to the left. Flagellum 1^-2 times the body length. Chloroplasts

Thompson : Fresh-water Algae 53

numerous, discoid. Length 33 mu, width 22 mu. The material collected
agrees well with the description of L. turbiniformis up to the description of
the tail. The tail of the individuals collected in Kansas is 9.9 mu long and
blunt at the apex, not just a sharp spine-like projection as figured and
described by Deflandre. The eyespot is at the anterior end nearly median
in position and slightly below the depression in which the flagellum is inserted.
Coll. April, May, June, July, 1937, boat slough at north end of Ohio street,
Lawrence; June, Chetopa creek, Altoona.

Lepocdnclis texta (Duj.) Lemm.

(PL XI, fig. 9)

Cells broadly oval, 52-60 mu long, 38 mu wide. Periplast strongly spirally
striated to the right. Flagellum 2-3 times the body length. Paramylum
grains numerous, cylindric or ring-shaped. Chloroplasts numerous, discoid.
The individuals collected measured 49.5-51 mu long, 39.6-42.9 mu wide.

Coll. June, July, 1937, Green's pond, Lawrence; June, Chetopa creek pool,
Altoona.

Lepocinclis truncata Da Cunha
(Conrad 1934)

(PI. XI, fig. 10)

Cells broadly ovoid to rhomboid with the anterior pole truncated and
flat and the posterior pole round-pointed, 40 mu long, 28 mu wide. The ma-
terial collected measured 46 mu long, 33 mu wide. Periplast delicately spirally
striated to the left. Chloroplasts numerous, discoid. Flagellum 1-1% times
body length. Two large thick paramylum rings are present.

Coll. June, 1937, Chetopa creek pool, Altoona.

Phacus longicauda (Ehrenb.) Duj.

(PI. XI, fig. 11)

Cells oval with a long attenuated tail, 85-144.4 mu long, 46-70 mu wide.
Periplast longitudinally striated. Flagellum shorter than the body length.
There is one large disk-shaped or ring-shaped paramylum body in each cell.

Coll. April, May, June, July, 1937, boat slough at north end of Ohio street,
Green's pond, Lawrence; Chetopa creek pool, Altoona.

Phacus longicauda (Ehrenb.) Duj. var, torta Lemm.

(PI. XI, fig. 12)

Similar to P. longicauda in all respects except that the individuals are
strongly twisted, like a propeller.

Coll. spring, 1937, boat slough at north end of Ohio street, Lawrence.

Phacus anacoelus Stokes

(PL XI, fig. 13)

Cells oval or roundish, inflated into five flanges with concave sides, 42.9 mu
long, 39.6 mu wide. Periplast longitudinally striated. Tail short. Flagellum
body length. 1-2 paramylum rings may be present.

Coll. July, 1937, Green's pond, Lawrence; Village creek, Chanute.

54 The University Science Bulletin

Phacus acuminata Stokes

(PI. XI, fig. 14)

Cells broadly egg-shaped or almost circular, 21-25 mil long, 14-25 mil wide.
Posterior end suddenly pointed. Periplast longitudinally striated. Median
fold on the dorsal surface comblike, nearly reaching the posterior end. Flagel-
lum body length. Two small round paramylum bodies are present.

Coll. April-July, 1937, boat slough at north end of Ohio street, Lawrence.

Other individuals seen in culture are illustrated in figure 20, plate XI. Their
dimensions lie near those of P. acuminata, they are of the same shape, varying
a little, and show no prominent ridge along the dorsal surface. However, from
the fact that they were all seen in the same culture as P. acuminata and that
they so nearly approximate P. acuminata in size and appearance, it is felt that
they are possible variants. The paramylum bodies vary in number and size.
The three individuals were chosen to figure as representative of the three de-
grees of variation that occurred in the culture. All forms were numerous, in-
cluding those that exactly fit the description of P. acuminata.

Figure 20 (a) 19.8 mu long, 9.9 mu wide.

Figure 20 (b) 22.8 mu long, 19 mu wide.

Figure 20 (c) 19.8 mu long, 13.2 mu wide.

Phacus pusilla Lemm.

(PI. XI, fig. 15)

Cells long-oval, 20 mu long, 7.5 mu wide, both sides with winglike infla-
tions. Anterior end concave, posterior end short pointed to almost round.
Periplast spirally striated. Flagellum one half body length. Two ringlike
paramylum bodies may be present.

Coll. July, 1937, stock pool in the sand hills, Burrton.

Phactis caudata Hubner

(PI. XI, fig. 16)

Cells long-oval, 45 mu long, 22.5 mu wide. Tail short, dorsal fold reaching
to the posterior end. Periplast longitudinally striated. Flagellum body length.
One large paramylum ring before the nucleus, a smaller one behind near the
tail.

Coll. May, 1937, boat slough at north end of Ohio street, Lawrence.

Phactis pyrum (Ehrenb.) Stein

(PL XI, fig. 17)

Cells o^^oid, posterior portion attenuated gradually into a long tail, 30-55
mu long, 13-15 mu wide. Periplast spirally striated to the left, striae very
strong. Flagellum body length. Two large or more smaller paramylum bodies,
disk-shaped, lie to the sides. Few large discoid chloroplasts are present.

Coll. April-July, 1937, boat slough at north end of Ohio street, Lawrence;
June, Village creek swamp, Chanute.

' Phacus triqueter (Ehrenb.) Duj.

(PI. XI, fig. IS)

Cells oval, 49-55 mu long, 33-35 mu wide. Tail short, sharp. Dorsal fold
comblike, reaching the posterior end. Periplast longitudinally striated. Flagel-
lum body length. One to two paramylum rings may be present.

Coll. June-July, 1937, Green's pond, Lawrence.

Thompson: Fresh-water Algae 55

Phacus Jtispidula (Eichw.) Lcmm.

(PI. XI, fig. 19)

Cells oval, with a short tubular flagellum opening and a short, blunt tail,
30-35 mu long, 18-33 mu wide. Periplast longitudinally striated. Striae finely
toothed with small spines. Flagellum body length. Paramylum bodies stick-
like to discoid.

Coll. July, 1937, Green's pond, Lawrence.

Phacus inflata sp. nov.

(PI. XI, fig. 21)

Cellula fere quadrata est, ex posteriore parte brevissima et obtuse gibbosa;
42.2 mu longa, 42.2 mu lata; membranum non manifesto striatum est; flagel-
lum par est longitudini corporis; chlorophorae multae et discoides sunt.

Cells almost quadratic or obscurely five-sided with a very short, blunt pos-
terior protuberance, 42.2 mu long, 42.2 mu wide, 14 mu thick. Periplast not
striated. Flagellum body length. Chloroplasts numerous and discoid.

Coll. June, 1937, Village creek, Chanute.

Crypotoglena pigra Ehrenb.

(PI. XII, fig. 2)

Cells ovate and somewhat compressed. Anterior pole broadly rounded,
posterior pole acute. Two laminate, longitudinal chloroplasts, one on each
side of the cell. Cells 11-15 mu long, 6-9.5 mu wide. Flagellum body length.

Coll. March-April, 1937, Jackman's pond. Green's pond, boat slough at
north end of Ohio street, Lawrence.

Trachelomonas oblonga Lem-m.

(PI. XII, fig. 3)

Lorica oval, 13-16 mu long, 11-12 mu wide, with or without a short collar.
Flagellum opening circular. Lorica yellow-brown to deep red-brown, some-
times covered with irregular deposits of iron compounds.

Coll. March-July, 1937, boat slough at north end of Ohio street, Green's
pond, Lawrence; July, stock pool in sand hills, Burrton.

Trachelomonas volvoaina Ehrenb.

(PL XII, fig. 1)

Lorica spherical to slightly ovoid, 7-21 mu in diameter, light to deep yellow-
brown, smooth. Flagellum opening circular, seldom with a collar. Flagellum
2-3 times the length of the lorica.

Coll. March-July, 1937, Green's pond, Lawrence; April, June, Village creek
swamp, Chanute.

Trachelo77ionas volvocinu Ehrenb. var. cervicula (Stokes) Lemm.

(PI. XII, fig. 4)

Lorica spherical, 23 mu in diameter, smooth, yellow-brown. Flagellum open-
ing circular with a tubular collar that projects inward as well as exteriorly.
Chloroplasts numerous and discoid. Flagellum 2-3 times diameter of the lorica.

Coll. July, 1937, Village creek swamp, Chanute.

56 The University Science Bulletin

Trachelomonas hispida (Perty) Stein var. cylindrica Klebs

(PI. XII, fig. 5)

Lorica cylindrical, thickly ornamented with very fine spinelike granules,
23-26.4 mu long, 14.8-16.5 mu wide. Flagellum opening collared with a toothed
crown. Flagellum 2-3 times lorica length.

Coll. July, 1937, stock pool in the sand hills, Burrton.

Trachelomonas his'pida (Perty) Stein

(PI. XII, fig. 6)

Lorica oval, thickly ornamented with very fine spines, 20 to 42 mu long,
15-26 mu wide. Flagellum opening with a short collar which may be toothed
and may be lacking. Chloroplasts 8-10, each with a double-layered pyrenoid.
Flagellum l%-2 times body length.

Coll. July, 1937, stock pool in the sand hills, Burrton; boat slough at north
end of Ohio street, Lawrence.

Trachelomonas Vermont! Deflandre (Pascher 1929)

(PI. XII, fig. 9)

Lorica in cross section four-sided, the angles rounded and the sides inflated
shghtly. Lorica from one side cylindrical to broadly ovoid, at the base ending
in a blunt, short point, smooth or lightly granular. Chloroplasts 8-12, parietal,
discoid and without pyrenoids. Flagellum IV2 times length of the lorica.

Coll. July, 1937, stock pool in the sand hills, Burrton.

Tranchelomonas armata (Ehrenb.) Stein

(PI. XII, fig. 7)

Lorica broadly oval, not punctate, ornamented at the posterior end with
a circle of long spines, 29-64 mu long, 31.3 mu wide. Flagellum opening thick-
ened or with a toothed collar. Flagellum twice the length of the lorica.

Coll. June, July, 1937, Green's pond, boat slough swamp at north end of
Ohio street, Lawrence.

Trachelomonas pidchra var. elongata Swirenko (Pascher 1929)

(PI. XII, fig. 8)

Lorica broadly ellipsoidal, brown, 35-39 mu long, 26-30 mu wide. Anterior
and posterior poles ornamented with seemingly irregularly placed, short, dull
spines. Chloroplasts numerous, discoid, up to 8 mu wide, without pj'renoids.
Flagellum three times the length of the lorica.

Coll. July, 1937, Green's pond, boat slough swamp at north end of Ohio
street, Lawrence.

Thompson: Fresh-water Algae 57

CHAROPHYTA

Family Characeae

subfamily nitelleae

Nitella opaca Ag.

(PI. XII, fig. 10)

Plant bodj' becomes a large, widespreading bush in quiet water. It is
dioecious, not encrusted with calcium and there is no gelatinous sheath around
the sex organs. The leaves are but once divided and the end segment is one-
celled. The corona on the oogonium is deciduous. It is made up of five two-
celled tips. The tips of the five spirally wound oogonial sheath cells spread
after the shedding of the corona. The oogonia may be one to three at a node.
The antheridia are always single at a node. The zygote with 6-7 spiral flanges
is 300-360 mu long, and almost as broad. The antheridiura is 464-496 mu in
diameter.

Coll. April, 1937, small creek pool V2 mile we.st of town, Altamont.

SUBFAMILY CHAREAE

Chara foetida A. Br.

(PI. XII, fig. 11)

Plant entirely encrusted except for the subulate tips of the leaves, monoeci-
ous, the oogonia and antheridia being borne at the same node with the oogonia
above the antheridia. Leaves 6-11 in a whirl at a node, 8 in a whirl in the
material collected. Stipules in two rows or series. Internodal cells twice as
many as there are leaves in the node above. Node cells or the blunt pro-
tuberances at the nodes of the internodal cells occur in the furrows between
the overhanging internodal cells on each side. The leaflets at a node on the
opposite side to the sex organs almost entirely undeveloped, scarcely longer
than broad. Antheridia 250-300 mu in diameter. Oogonia 750-800 mu long,
450 to 500 mu wide, with 12-15 spirals.

Coll. April, 1935. Deer creek, Garnett; June, 1937, Lost canyon side hill
swamp and spring, Neodesha.

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Taylor. "VVm. Randolph. (1934) The Fresh-ivater Algae of Newfoundland I.
Mich. Acad. Sci., Aiis and Letters. 19: pp. 217-278.

(1935) The Fresh-water Algae of Newfoundland II. Mich. Acad. Sci.,

Arts and Letters. 20: pp. 185-230.

TiLDEN, Josephine. Minnesota Algae Vol. I. Minn. Univ., Minneapolis, 1910.
Walton, L. B. (1915) A review of the described species of the order Eugle-

noidina Bloch. Ohio State Univ. Bull. 19: 343-459.
West, G. S., and Fritsch, F. E. British Fresh-water Algae. Cambridge Univ.

Press, 1927.
WoLLE, Francis (1887) Fresh-ivater Algae of the United States. Comenius

Press, Bethlehem, Pa.

60 The University Science Bulletin

PLATE I

1. Chroococcus twgidus. X 166.

2. Chroococcus caldariorum. X 166.

3. Chroococcus limneticus. X 166.

4. Gleocapsa magma. X 166.

5. Gleocapsa viellea. X 166.

6. Gleocapsa aeruginosa. X 166.

7. Microcystis marginata. X 166.

8. Merismopedia punctata. X 333.

9. Merismopedia elcgans. X 333.

10. Merismopedia elegansf (See text.) X 125.

11. Eucapsis alpina. X 333.

12. Aphanothece microscopica.

(a) Colony. X 83.
(6) Single cell. X 333.

13. Pleiirocapsa minor. X 166.

14. Merismopedia angularis sp. nov. X 210.

(6) Side view.

15. Chnmaesiphon cylindricus. X 666.

16. Oscillatroroia curviceps. X 333.

17. Chamaesiphon gracilis. X 666.

18. OsciUatoria sancta. X 166.

19. Oscillatorin simplicissima . X 333.

20. Spirulina major. X 333.

21. OsciUatoria guttidata. X 333.

22. OsciUatoria jcnensis. X 166.

23. Arthrospira jenneri. X 333.

24. Phormidium tenue. X 333.

25. Lynghya Birgei. X 166.

26. Microcolvus vaginatus. X 333.

27. Lyngbya aerugineo-caerulea.

(a) Spore formation. X 333.
(6) Vegetative filament. X 333.

1

Thompson: Fresh-water Algae

61

PLATE I

62 The University Science Bulletin

PLATE II

1. Nostoc verrucosum.

(a) Colony. X 333.

(b) Single filament. X 333.

2. Nostoc juscescens var. mixta. X 333.

3. Nostoc coeruleum. X 333.

4. Anahaena calijornica. X 333.

5. Nostoc viuscorum. X 333.

6. Cylindrospermum viaius. X 333.

7. Cylindrospermmn minutissimum. X 333.

8. Rivularia planctonica. X 333.

9. Calothrix Braunii. X 333.

10. Rivularia compacta. X 333.

11. Plectonema Golenkiniana. X 666.

12. Scytonema myochrous. X 333.

13. Porphyndium crventum.

(a) Cells. X166.

(6) After Lewis and Zirkle, 1920. X 200.

14. Botrydium granvlatum.

(a) Vegetative thallus. X 16.6.
(6) Thallus breaking up into hypnospores.
X 16.6.

(c) Variously shaped hj-pnospores. X 63.

15. Synura uvella.

(a) Mature colony. X 166.

(b) Small young colony. X 166.

(c) Single individual. X 333.

16. Mallonionas toit.mrotn. X 333.

17. Chromulina ovalis. X 333.

18. Cryptomonas ovata. X 1'70.

19. Tribonema bombycinum. X 166.

20. Phaeothainnion Borzianuvi.

Palmella stage. X 166.

Thompson: Fresh-water Algae

63

PLATE II

64 The University Science Bulletin

PLATE III

1. Chlamydomonas intermedia.

(a) Motile individual. X 333.

(b) Palmella stage. X 333.

2. Chlorogonium enchlorum.

(a) Adult. X333.

(b) Zoospore production. X 333.

(c) Zoospores escaping. X 333.

3. Chlorogonium spirale. X 333.

4. Wislouchiella planctonica.

(a) Front view. X 250.

(b) Side view. X 250.

(c) Top view. X 250.

5. Carteria cordiformis. X 250.

6. Chlamydomonas variabilis. X 333.

7. Phacotus lenticularis.

(a) Front and side views. X 333.

(b) Palmella stage. X 333.

(c) Zoospore production. X 333.

8. Gonium sociale.

(a) Top view. X 166.

(b) See text. X 166.

9. Gonium jormosum. X 166.

10. Pandorina 7norum.

(a) Vegetative colony. X 166.

(b) Sexual colony. X 166.

11. Eudorina elegans. X 166.

1

Thompson: Fresh-water Algae
PLATE III

65

66 The University Science Bulletin

PLATE IV

1. Spondyloviorum quaternarium. X 292.

2. Sphaerella lacustris. X 166.

(a) Motile individual.

(b) Akinete.

(c) Microgamete.

(d) Aplanospore.

(e) Akinete germinating by zoospores.

3. Tetraspora gelatinosa. X 250.

4. Ourococcus bicaudatus. X 333.

5. Tetraspora cylindrica. X 250.

6. Coccomyxa dispar. X 333.

7. Nannochloris bacillaris. X 333.

8. Chlorosarcina consociata. X 166.

9. Ulothrix tenuissima. X 166.

10. Gleotila prologenita. X 166.

11. Hormidium flaccidum. X 166.

12. Stichococcus subtilis. X 166.

13. Microspora tumidula. X 166.

14. Microspora stagnorum. X 166.

15. Geminella ordinata. X 166.

16. Cylindrocapsa geminella. X 166.

17. Chaetovhora elegans. X 166.

18. Draparnaldia Ravenelii. X 16.6.

19. Chaetophora incrassata. X 166.

20. Draparnaldia plumosa. X 16-6.

Thompson : Fresh-water Algae

67

PLATE IV

68 The University Science Bulletin

PLATE V

1. Microthamnion Kuetzingianum. X 166.

2. Aphanochaeie repens. X 83.

3. Ulvella involvens. X 166.

4. Coleochaete scutata. X 33.

5. Rhizoclonium cnspum. X 16.6.

6. Chaetopeltis orhiculaiis. X 8.

7. Cladophora glomerata. X 33.

8. Pithophora kewensis. X 20.

9. Basicladia chelonum. X41.

10, Oedogonium globosum. X 166.

11. Oedogonium crispum. X 166.

Thompson : Fresh-water Algae

69

PLATE V

70 The University Science Bulletin

PLATE VI

1. Oedogonium cyathigerum. X 83.

2. Bv^hochaete varians. X 83.

3. Schizomeris Leibleinii. X41.

(a) Portions at intervals of a thallus.

ib) (c) Terminal and intercalary zoosporangia.

id) Different apical cells.

4. Chlorococcum humicola. X 333.

5. Chlorococcum injusionum. X 333.

6. Schizogonium murale. X 166.

7. C hlorochytrium lemnae. X 93.

8. Golenkinia radiata. X 166.

9. Characium omithocephalum. X 750.

10. Characium angustum. X 750.

11. Characium Naegelii. X 750.

12. Characium Braunii. X 750.

13. Pediastrum Boryanum. J

(a) Typical coenobium. X 125.
(t>) Four-celled coenobium. X 166.

14. Pediastrum Boryanum, var. brevicorne. X 166.

15. Pediastrum duplex var. comutum. X 83.

16. Pediastrum duplex var. reticulatum. X 166.

17. Pediastrum, tetras. X 166.

18. Pediastrum duplex var. gracillimum. X 125.

19. Pediastrum, Boryanum var. longicome. X 166.

Thompson : Fresh-water Algae

71

PLATE VI

72 The University Science Bulletin

PLATE VII

1. Pediastrum duplex var. clathratum. X 83.

2. Pediastnim clathratum. X 333.

3. Pediastrum simplex var. radians. X 125.

4. Pediastrum clathratum var. duodenarium. X 166

5. Coelastrum sphaericiim. X 166.

6. Coelastrum microporum. X 166.

7. Coelastrum reticulatum. X 166.

8. Chlorella conductrix.

(a) Single cell. X 333.
(6) Habit. X 83.

9. Westella botryoides. X 166.

10. Dictyosphaerium pulchellum. X 333.

11. Trochiscia pachyderma. X 166.

12. Excentrosphaera viridis. X 166.

13. Schroederia setigera. X 333.

14. Oocystis parva. X 333.

15. Oocystis elUptica. X 103.

16. Diinorphococcus lunatus. X 333.

17. Closteridium lunula? X 333.

18. Ankistrodesmus falcatus var. mirabile. X 333.

19. Ankistrodesmus falcatus var. acicularis. X 333.

4
1

Thompson : Fresh-water Algae

73

PLATE VII

74 The University Science Bulletin

PLATE VIII

1. Sclenastrum gracile. X 166.

2. Kirchneriella contorta. X 333.

3. Kirchneriella lunaris. X 333.

4. Tetraedron caudatuvi. X 250.
5 Tetraedron minimum. X 250.

6. Tetraedron punctulatimi. X 250.

7. Tetraedron trilohatum. X 166.

8. Tetraedron pentaedricum. X 166.

9. Tetraedron trigonum var. minor. X 166.

10. Tetraedron tumididum. X 166.

11. Tetraedron proteiforme. X 83.

12. Scenedesmus obliquus. X 333.

13. Scenedesmus dimorphus. X 333.

14. Scenedesmus bijuga. X 333.

15. Scenedesmus quadricauda. X 333.

16. Scenedesmus abundans. X 333.

17. Crucigenia rectangularis. X 333.

18. Scenedesmus denticulatus. X 308.

19. Actinastrum, gracillimum. X 375.

20. Actinastrum Hantzschii var. javanicum.

X375.

21. Tetras-trum apiculatum. X 333.

22. Tetrastrum elegans. X 333.

23. Crucigenia tetrapedia. X 333.

24. Micractinium ptisHlum. X 250.

25. Errerella bornhemieyisis. X 250.

Thompson: Fresh-water Algae

75

PLATE VIII

76 The University Science Bulletin

PLATE IX

1. Vaucheria geminota. X 63

2. Vaucheria sessilis. X 63.

3. Zygnema insigne. X 83.

4. Spirogyra protecta. X41.

5. Spirogyra malmeana. X41.

6. Spirogyra punctiformis. X31.

7. Spirotaenia parvula. X 333.

8. i?o|/a obtusa. X 125.

9. Netrium digitus. X 63.

10. Closterium subulaium. X 63.

11. Closterium acerosutn. X 29.

12. Closterium moniliforme. X 29.

13. Spirogyra decimina. X 166.

14. Pleurotacnium Ehrenbergii. X 63.

15. Penium curcubitum. X 166.

16. Cosmarium reniforme. X 166.

17. Cosmarium polymorphum. X 166.

18. Cosmarium punctulatum. X 166.

19. Cosmarium, circidare. X 166.

20. Cosmarium, pseudobroomei. X 166.

21. Cosmarium Schliephackeanum. X 83.

22. Euastrum, verrucosum. X 125.

Thompson : Fresh-water Algae

77

PLATE IX

78 The University Science Bulletin

PLATE X

1. Cosmarium polygonum. X 166.

(For figures lb and Ic see text.)

2. Cosmarium granatura. X 166.

(For figures 2&, 2c, see text.)

3. Cosmarium impressulum. X 166.

4. Cosmarium sexpapillosiim sp. nov. X 166.

5. Cosmarium. Nymannianum,? X 166.

6. Cosmarium humile var. lacustre. X 166.

7. Cosmarium crenaLum. X 166.

8. Micrasterias am,ericana. X 63.

9. Staurastrum, paradoxum. X 166.

10. Glenodinium oculatum. X 250.

11. Glenodinium pulvisculus. X 250.

12. Euglena intermedia. X 125.

13. Euglena spirogyra. X 166.

14. Euglena acus. X 166.

15. Euglena rubra. X 125.

16. Euglena truncata var. baculifera var. nov.

X125.

17. Euglena alata sp. nov. X 125.

18. Euglena deses. X 125.

19. Euglena spiroides. X 125.

20. Euglena Grisoli. X 125.

21. Euglena jusca. X 125.

22. Euglena genicidata. X 125.

23. Euglena iripteris. X 125.

24. Euglena torta. X 125.

25. Euglena viridis. X 125.

26. Euglena proxima. X 125.

27. Euglena variabilis. X 125.

28. Euglena acutissima. X 125.

29. Euglena flava. X 125.

Thompson: Fresh-water Algae

79

PLATE X

80 The University Science Bulletin

PLATE XI

1. Lepocinclis ovum. X 250.

2. Lepocinclis ovum var globula. X 250.

3. Lepocinclis ovum var. palatina. X 250.

4. Lepocinclis Butschlii var. angustata. X 250.

5. Lepocinclis acicularis. X 250.

6. Lepocinclis Steinii. X 250.

7. Lepocinclis turhinijormis. X 250.

8. Lepocinclis jusijormis. X 250.

9. Lepocinclis texta. X 250.

10. Lepocinclis tnincala. X 250.

11. Phacus longicauda. X 250.

12. Phacus longicauda var. ^oria. X 250.

13. Phacus anacoelus. X 250.

14. Phacus acuminata. X 250.

15. Phacus pusilla. X 250.

16. Phacus caudata. X 666.

17. Phacus pyrum. X 666.

18. Phacus triqueter. X 250.

19. Phacus hispidula. X 250.

20. Phacris acuminata 1 X-250.

(For figures 20o, h and c, see text.)

21. Phacus inflata sp. nov. X 250.

Thompson : Fresh-water Algae

81

6-^141

82 The University Science Bulletin

PLATE XII

1. Trachelomonas volvocina. X 250.

2. Cryptoglena pigra. X 666.

3. Trachelomonas ohlonga. X 666.

4. Trachelomonas volvocina var. cervicula. X 250.

5. Trachelomonas hispida var. cylindrica. X 250.

6. Trachelomonas hispida. X 250.

7. Trachelomonas armata. X 250.

8. Trachelomonas pidchra var. elongata. X 250.

9. Trachelomonas Verm,onti. X 250.

10. Nitella opaca.

(a) Antheridum. X 33.

ib) Leaf tip. X 33.

(c) Oogonium without corona. X 33.

id) Corona. X 166.

(e) Zygote. X 33.

(/) Habit sketch. X -12.

11. Chara joetida.

(a) Portion of a thaUus bearing an oogonium
and an antheridium. X 33.

(6) Leaf tip. X 33.

(c) Portion of stem showing stipules and cor-
tical outgrowths. X 16.6.

Thompson : Fresh-water Algae
PLATE XII

83

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 2

Effects of Certain Alkaloids on the Growth of Aspergillus
niger and Rhizopus nigricans

JAMES C. BATES,
Department of Botany, University of Kansas

Abstract: A study was made on the effect of caffeine, caffeine citrate, and
the sulphates of strychnine and quinidine on the growth and sporulation of
Aspergillus niger van Tieghem and Rhizopus nigricans Ehrhart on Coon's nu-
trient solution. The results showed that, in general, the rate and amount of
growth was increased by strychnine and quinidine and decreased by caffeine
and caffeine citrate. Sporulation was retarded or completely suppressed by all
of the above alkaloids except strychnine.

A second study was made to determine the ability of Aspergillus niger to
assimilate strychnine-carbon and nitrogen. The fungus was grown on different
nutrient solutions in which the only carbon and nitrogen containing com-
pounds present were as follows: strychnine and KNO3; dextrose and strych-
nine; dextrose and KNO3; dextrose, KNO.3 and strychnine. The results of
this experiment showed that strychnine-carbon, when the only carbon com-
pound present, is unavailable and that strychnine-nitrogen is utilized to a
slight extent in the absence of other nitrogen-containing compounds.

A study was then made to determine the ability of Aspergillus niger to
assimilate strychnine-carbon and nitrogen in the presence of dextrose and po-
tassium nitrate. The amount of growth obtained on nutrient solutions with
varying ratios of dextrose and KNO.s and equal amounts of stiychnine was
compared with that obtained on control solutions without the strychnine.
The results showed that the increase in rate and amount of growth due to the
presence of strychnine was as great in solutions with a low, as in those with a
high, dextrose/KN03 ratio. Moreover, quantitative analysis of the culture
solutions showed that their stiychnine content remained practically unchanged.
The results of these experiments indicate that strychnine has an intense stimu-
lating action on growth and very little, if any, nutritive value.

INTRODUCTION

IN undertaking a study of the effects of alkaloids on the growth
of fungi, one naturally wonders as to the origin and role of these
substances within the plants which produce them.

The alkaloids are rather limited in their distribution in the plant
kingdom, occurring, for the most part, in the Leguminosae, Papa-

(8.5)

86 The University Science Bulletin

veraceae, Ranunculaceae, Rubiaceae, and Solanaceae. They may
occur in solution in the cell sap in the young parenchyma or may be
stored in older tissue in the solid state. They are generally found
in the seeds and fruits, but some occur in the leaves, stems and roots
(Hass and Hill, 13).

The origin of the alkaloids in the plant is not definitely known.
Some consider that they arise in the formation of proteins, while
others believe that they are formed in the disintegration of the pro-
teins. They are thought to be nitrogenous waste products by most
investigators.

As to the role of the alkaloids within the plants which produce
them, there have been differences in opinion among the investigators.
Some have ascribed to these substances a protective function against
the attack of animals. The literature on this subject indicates that
the protective function, when performed, is only an incidental one.

Some have ascribed to the alkaloids a nutritive value clue espe-
cially to the nitrogen contained in their molecules. A part of the
evidence bearing on this subject comes from investigations of the
localization of alkaloids, their quantitative variations, mode of for-
mation, migration during the course of vegetation and of seed germi-
nation, variations provoked under the influence of different external
factors, and grafting experiments.

The literature of the above-mentioned investigations has been
contributed by numerous investigators and summarized by Goris
(12). From the following brief summary of these investigations,
taken from Goris, one may obtain some idea as to the general nature
of this work and its bearing on the role of alkaloids.

The localization of alkaloids has been studied by microchemical
reactions produced within the tissues of the plant under the influ-
ence of various reagents. Some of these reagents form insoluble
precipitates with the alkaloids while others give beautiful color re-
actions which are, in many cases, of but short duration. These tests
give but little information concerning the actual amount of alkaloid
present.

The alkaloids have been found freciuently and in considerable
amount in the latex vessels of the Papaveraceae and Fumariaceae,
the secreting cells of Conium maculatutn L. and associated with
raphides in the Amaryllidaceae.

In general, the alkaloids are found in regions where cellular ac-
tivity is greatest.

From his study of the work on localization, Goris concludes that

Bates: Effects of Alkaloids on Growth 87

the alkaloids are translocated, to a great extent, to external tissues
of the stem which are later sloughed off. Goris also accepts the
presence of alkaloids largely in the secretory tissue as evidence that
they are not reserve compounds. Several hypotheses, based on
similarities and differences between structural formulae of alka-
loid molecules and those of various other compounds found in plants,
have been advanced to explain the mode of formation of alkaloids.
The most commonly accepted hypothesis seems to be that of Winter-
stein and Trier (34), who consider the alkaloids as waste products
arising from the metabolism of albuminous material. The origin
of the purine bases from the decomposition of nucleoproteins is
commonly accepted. It is interesting to note the frequency with
which certain cyclic nitrogenous groups are common to both al-
kaloids and aminoacids, for example, proline and nicotine both
contain the pyrrolidine ring while the indole ring is common to
tryptophane and brucine.

Cimician and Ravenna (3) injected various nitrogen compounds
into plants which normally contain alkaloids. The alkaloid con-
tent of these plants was determined after a time and compared to
controls which had not been injected.

The nicotine content of the tobacco plant injected with aspara-
gine was greater than that of the control plant.

Additional evidence bearing on the nutritive value of alkaloids
comes from experiments on the effects of these substances on seed
germination and the growth of different plants. This problem deal-
ing with the effects of alkaloids on the growth of fungi warrants a
brief review of the literature on the previous work on this subject.

REVIEW OF LITERATURE

According to Wood, Remington, and Sadtler {36) strychnine sul-
phate is used in practical medicine in doses of one fortieth to one
twentieth grain (0.0016 to 0.003 gm.). One quarter of a grain has
been known to produce death within a few hours.

Quinidine sulphate has a similar physiological action to that of
quinine. Antimalarial dose, one gram.

Caffeine is a valuable remedy in practical medicine as a cerebral
and cardiac stimulant and as a diuretic. Dose, from three to eight
grains (0.2 to 0.5 gm.).

Caffeine citrate is possessed of the therapeutic properties of caf-
feine. Dose, one teaspoonful (3.9 gm.).

Overton {36) observed that caffeine and strychnine quickly pene-
trated the protoplasm of plant and animal cells.

88 The University Science Bulletin

Marcacci (19) found that atropine and morphine produced a
favorable action on lactic acid fermentation which was retarded by
quinine, veratrine, cinchonamine, and to a still greater extent by-
strychnine. All of the above alkaloids produced a favorable action
on alcoholic fermentation except cinchonamine and quinine.

Considerable work has been done on the effect of alkaloids on
the growth of the higher plants. The experimental plants were
grown in nutrient solutions, sand, or soil to which the alkaloid was
added. The controls were grown in nutrient solutions without the
alkaloids or in sand or soil sprinkled with pure water instead of the
alkaloid solution. A large number of different plants and alkaloids
have been used.

A number of investigators, Sawa (29), Lutz {16), Strake (30),
Weyl (33), Marcet(^O), Bouchardat (^), Wolf and Knop(35),
Marcacci (19), Otto (^5), Detoni(5), and DeVarigny (5), found
that the alkaloids produce a toxic or unfavorable action on the
growth of the higher plants. Princeps {27) found that plant ex-
tracts of alkaloid-containing plants produced an injurious action
on the same plants from which they were obtained.

It has been observed by Strake {30) , Marcet {20) , Bouchardat
{2), Wolf and Knop (35) and Marcacci [19) that the toxic action
varies with different alkaloids.

DeVarigny (5) , Bouchardat (;?) , and Otto (.55) found that the
toxic action of the alkaloids varies inversely with the fertility of
the soil in which the experimental plants are grown. Goris {12)
attributes the increased resistance of the plant growing in fertile soil
to the greater abundance of nitrate which enables the plant to
utilize the alkaloid. It seems to the writer that the colloidal action
of the soil may also play a part in decreasing the injurious action
of the alkaloids.

Reveil {£8) found that plants sprinkled with atropine solutions
grew more vigorously than the controls. He identified the alkaloid
in the experimental plants and concluded from his work that the
alkaloid assimilated disappeared rapidly during the course of
growth. His experiments were repeated by Marcacci {19) and De-
Varigny {9), who were unable to confirm them. Wolf and Knop
{35) were unable to identify the alkaloid in the experimental plants
with which they worked.

Goeppert {11) observed that seedlings of oats, wheat, peas, and
cress sprinkled with infusions of belladonna developed as well as
those sprinkled with pure water. DeVarigny {9} obtained experi-
mental plants superior to the controls by adding atropine solution to

Bates: Effects of Alkaloids on Growth 89

the soil. When sand was used instead of soil the experimental plants
were inferior to the controls. Marcacci {19) observed that Lemna
minor is uninjured by morphine and atropine. Bokorny {1) states
that the higher plants are very little susceptible to caffeine.

DeToni (8) grew Coix lacryma in nutrient solutions which con-
tained strychnine sulphate as the only nitrogen-containing com-
pound, but with only partial results.

Lutz {15) germinated seeds in washed and sterilized sand mois-
tened with a nutritive solution containing an alkaloid. He found
that during the development of the seedling there was a loss instead
of a gain in nitrogen. He concluded that the alkaloids when used
alone as a source of nitrogen were unassimilable.

Marcacci {19) observed that the roots are the most susceptible
part of the plant to the injurious action of alkaloids.

Marcacci {19) studied the influence of different alkaloid solutions,
which varied in concentration from 0.1 to 0.005 percent, on the ger-
mination of different seeds. He observed a toxic action which varied
with the nature of the alkaloid.

Mirande {22) obtained good germination of hemp seeds when
treated with valerianate of quinine in a dilution of 1-2500, but the
seedlings were inferior to the controls.

DeVarigny {9) studied the action of 1-500 and 1-1500 solutions
of atropine on the germination of different seeds. According to the
results obtained, the seeds could be divided into three categories:
those for which atropine was unfavorable, indifferent, or favorable.

Cornevin (7) studied the action of different alkaloids on the ger-
mination of different seeds. He found that in some cases the germi-
nation of the seeds was retarded by the alkaloid, in some cases it
was unaffected, and in others germination was advanced with a
greater percentage of the seeds germinating.

Mosso {23) accurately determined the doses of different alkaloids
necessary to produce a stimulating action on germination. He pre-
pared solutions with concentrations varying from 2 to 0.0001 per-
cent. He germinated seeds of Phaseolus inultijiorus Willd. on cotton
placed in the bottom of a vessel and saturated with the different
alkaloid solutions. He found the optimum concentration for each
alkaloid which produced seedlings far superior to the controls. The
growth of the seedlings decreased as the concentration of alkaloid
was increased above, or decreased below, the optimum value. The
optimum concentrations for the different alkaloids are: chlorohy-
drate of morphine, 0.001 percent; nicotine solutions, 0.01 percent;

90 The University Science Bulletin

strychnine solutions, 0.005 percent; cocaine solutions, 0.01-0.001 per-
cent; atropine solutions, 0.0005 to 0.0001 percent; and in caffeine
salicylate solutions all seedlings were inferior to the controls. The
negative results obtained in caffeine solutions were attributed to an-
tifermentative actions of salicylic acid.

These experiments show that in high concentrations the alkaloids
have a toxic action and in weak doses an exciting action on seed
germination. The concentrations which are injurious or beneficial
vary with each alkaloid and with each plant.

Bokorny (1) found from 0.01 to 0.1 percent solutions of strych-
nine injurious to algae and small water animals. He states that
caffeine is a weak poison for algae and infusoria.

Comere (6) grew Ulothrix subtilis and Spirogyra crassa Kutz
in a nutrient solution devoid of nitrogen compounds and to which
he progressively added alkaloid salts. Morphine, atropine, and
cocaine were directly assimilated in a decreasing order. The salts
of quinine were also assimilated while those of strychnine were toxic.
It recjuired a concentration of 0.002 percent to kill Spirogyra and
0.008 percent to kill Ulothrix.

Nobecourt (34) cultivated Botrytis cinerea Pers. on Raulins liquid
containing varying amounts of nicotine, sulphates of atropine and
quinine, and aconitine. He found that nicotine sulphate in a con-
centration of 2%ooo and atropine sulphate in a concentration of
2%ooo did not hinder the growth of this fungus. Growth was not
retarded by a concentration of ^9i(ioo of quinine sulphate; but re-
sulted in little thalli with few conidiophores by a concentration of
Wiaao and completely inhibited by a concentration of ^%ooo. Growth
was visibly hindered by aconitine in a concentration of 74ooo, and
greatly reduced by a concentration of "fiooo. A concentration of
^%ooo did not prevent the germination of spores, which was com-
pletely inhibited by a concentration of ^%ooo.

Yasuda (37) found that the growi:,h of Penicillium glaucum, As-
pergillus niger, Botrytis cinerea and Mucor stolonifer was increased
by the addition of the hydrochlorides of cocaine, quinine, cincho-
nine, morphine, codine, and strychnine to Richards nutrient solu-
tion. The alkaloids were used in concentrations varying from 0.2
to 2 percent. As the concentration of alkaloid was increased, the
conidiophores and sporangiophores became thinner and shorter.
Conidium and sporangium formation was entirely suppressed and
replaced by clamydospores when the optimum concentration for
fungus-vegGtation was suri^assed. The weakest alkaloid for the

Bates: Effects of Alkaloids on Growth 91

fungi under consideration was the hydrochloride of morphine, while
the strongest was that of cocaine. The fungi listed in order of their
decreasing resistance to alkaloids are: Penicillium glaucam, As-
pergillus niger, Botrytis cinerea, and Mucor stolonifer.

Ehrlich (10) grew different fungi on a mineral nutrient solution
to which he added different alkaloids as the only nitrogen-containing
compounds and ethyl alcohol or invert sugar as a source of carbon
in concentrations of 0.2 and 2 percent, respectively. A control series
without the alkaloid was run at the same time.

The fungi used were: Oidium lactis, Aspergillus niger, Penicillium
glaucum, Willia anomala, Pichia farinosa, a mixed culture obtained
by exposing the culture solution to the air, and an unknown species
of wine yeast. The alkaloids used were: pyridine, piperidine bitar-
trate, coniine, nicotine, cinchonine, quinine, brucine, cocaine and
morphine.

Ehrlich determined the amount of fungus dry weight produced
and nitrogen content of the fungus substance at the end of a growing
period, which varied from 3 to 12 months.

He obtained only a small amount of growth, which was least in
the yeast cultures and greatest in the mixed cultures. A small
amount of nitrogen corresponding to the amount of growth was
recovered from the fungus dry substance. He concluded that this
nitrogen was obtained by the organism from the decomposition of
the alkaloid. This decomposition was greater in the mixed cultures
under the united action of several organisms. He also obtained
the aromatic odor of ester compounds in some cases, and traces
of ammonia with Nessler's reagent in others. The checking of
growth which he obtained was attributed to the poisonous action of
the decomposition products of the alkaloids.

Lutz (16) grew Aspergillus niger and Penicillium glaucum on
different alkaloid solutions and control solutions with the same
elemental composition as that of Raulin's solution. The relative
amounts of fungus growth in grams of dry weight produced on the
different solutions were used as criteria of the nutritive value of
the solutions.

From the results of this investigation Lutz concluded that the
alkaloids when used alone as a source of nitrogen were not assimi-
lated by fungi, but were assimilated in- considerable quantities in
the presence of nitrate or ammonia-nitrogen.

In a later work, Lutz {17) set up an experiment to determine
whether fungi could use alkaloids alone as a source of nitrogen after
growth had been started on ammonium nitrate and to make quanti-

92 The University Science Bulletin

tative determinations of the amount of alkaloids utilized in both
the presence and absence of ammonium nitrate.

The fungi used were Aspergillus niger, A. repens, and Penicillimn
glaucum. The alkaloids used were the hydrochlorides of cocaine,
morphine and quinine.

He found some utilization of alkaloids from solutions whose nitro-
gen was only in the form of alkaloid-nitrogen. This he attributes
to traces of ammonium nitrate absorbed from Raulin's solution be-
fore replacement by the alkaloid solution and which could not be
washed from the mycelium. The amount of growth and of alkaloid
utilized were considerably greater in the alkaloid solutions contain-
ing ammonium nitrate. He reports an absorption of 0.947 gm. of
chlorohydrate of cocaine and 0.400 gm. of chlorohydrate of quinine
from 50 cc. of culture solution in 26 days.

Lutz suggests that the alkaloids are transformed into albumins in
the presence of ammonium nitrate. He does not consider them as
either reserve compounds or waste products in the strict sense of the
word, but as intermediate compounds between the albumins and
mineral nitrogen compounds.

From the literature reviewed on the alkaloids it appears that
these compounds when used in high dilution have an accelerating
action on growth and when used in high concentrations a toxic
action. The salts of the alkaloids seem to be less toxic than the
pure alkaloids. The intensity of action varies with different alka-
loids and wnth different experimental plants. In the higher plants
the roots and root hairs seem to be the most sensitive part of the
plant to the toxic action. The fungi seem to be least susceptible to
the toxic action of the alkaloids. In man\^ cases the fungi seem to
be benefited by high concentrations of these substances, while the
algae and higher plants are killed by very dilute solutions. It also
appears from the work of Ehrlick (1. c.) that some of the fungi are
able to assimilate small quantities of alkaloids in the absence of other
nitrogen-containing compounds. According to Lutz (1. c.) the alka-
loids may be assimilated in large quantities in the presence of nitrate
or ammonia nitrogen. Klotz (74) has shown that the nitrogen con-
tent of the fungus varies with the nitrogen and carbon sources of
the medium, the length of incubation, rate of growth, and the H-ion
concentration of the medium. A more acid medium prevents autol-
ysis and thereby tends to increase the nitrogen content of the fun-
gus. A rapid growth is accompanied by a lower percent of nitrogen
in the fungus. It follows from this that any substance which

Bates: Effects of Alkaloids on Growth 93

accelerates the rate of growth, if only by a purely stimulating action,
would effect a more efficient utilization of the nitrogen compounds
in the production of fungus dry weight.

With these results of previous investigators in mind the writer
has undertaken a study of the effect of caffeine, quinidine, and
strychnine on the growth of Rhizopus nigricans and Aspergillus
niger.

EXPERIMENTAL EFFECTS OF ALKALOIDS ON
GROWTH AND SPORULATION

Material and Method

The effect on growth was determined by growing a fungus on
100 c.c. portions of a nutrient medium containing varying amounts
of the alkaloid under investigation and making dry weight deter-
minations at regular intervals following inoculation. The relative
time required from inoculation until the first appearance of visible
mycelium and sporangia was used as a measure of the effect of the
alkaloid on germination and sporulation.

The fungi used, Rhizopus nigricans Ehrhart and Aspergillus
niger van Tieghem, were obtained from the stock cultures of the
Department of Botany, University of Kansas. These organisms
show considerable difference in their ability to use nitrate-nitrogen,
it being readily assimilated by Aspergillus niger, but only slightly
available to Rhizopus nigricans.

Coon's solution, as described by Young and Bennett (38), was
modified by increasing the sugar content to 2 percent, and used for
a culture medium to avoid difficulties of chemical analysis and
interpretation of results which would arise from using a culture
medium containing complex organic-nitrogen compounds.

Nutrient Solution

MsSOi 1 .23 grams

KH0PO4 2.72 grams

KN0.5 2.02 grams

Dextrose 20 . 00 grams

Distilled water 1000 c.c.

The above nutrient solution yields a good growth of Aspergillus
niger, but only a very scant growth of Rhizopus nigricans is ob-
tained unless the solution is reinforced with a more available form
of nitrogen.

The following alkaloids were used: sulphates of strychnine and
quinidine, caffeine citrate, and caffeine base. The alkaloids were

94 The University Science Bulletin

obtained from the Merck Chemical Company, weighed out in the
proper amounts and used without previous drying or purification.
Alkaloid solutions of varying concentrations were prepared by
adding the alkaloid in different amounts to 100 c.c. portions of the
nutrient solution. These 100 c.c. portions were placed in 125 c.c.
Erlenmeyer flasks, plugged with cotton, and sterilized in an Arnold
steam sterilizer.

A heavy suspension of the spores was obtained by growing the
fungus on potato dextrose agar and suspending the spores in dis-
tilled water. A drop of the heavy spore suspension was added to
each flask with a looped inoculating needle or dropped from a bur-
ette. All flasks were inoculated at the same time and from the
same spore suspension.

Dry-weight determinations were made at the end of a twenty-
three-day growing period in the experiments with Rhizopus nigricans
and at regular seven-day intervals following inoculation in the ex-
periments with Aspergillus niger. The mycelium was obtained by
filtering the solution through Gooch crucibles which previously had
been partially filled with asbestos fibers, dried to constant weight
in a desiccator and weighed. The mycelium was washed by running
distilled water through the filter. The filtering was hastened by
using a filter flask connected to a vacuum pump. The mycelium
obtained from five culture flasks was used for each dry-weight
determination in the experiments with Rhizopus nigricans. In the
experiments with Aspergillus niger the mycelium from two culture
flasks was used for each determination. The crucibles with my-
celium and spores were dried and reweighed as describe'd above.
The increase in weight represents the amount of fungus growth.

Results

Table I shows the effect of different concentrations of strychnine
sulphate, caffeine, caffeine citrate, and quinidine sulphate on the
growth of Rhizopus nigricans at room temperature for a twenty-
three-day growing period. Column I shows the alkaloid concen-
trations of the different nutrient solutions. Columns II, III, IV, and
V show the amount of growth obtained in the strychnine sulphate,
caffeine, caffeine citrate, and quinidine sulphate solutions, respec-
tively. The small amount of growth obtained was due to the in-
ability of Rhizopus nigricans to assimilate either the alkaloid or
nitrate-nitrogen as was later demonstrated by adding peptone to
the culture solution, in which case a vigorous growth was obtained.

The table shows that the amount of growth was greater in the

1

Bates: Effects of Alkaloids on Growth 95

nutrient solution to which strychnine sulphate was added than in the
nutrient solution alone; the amount of growth varying directly with
the concentration of strychnine sulphate and inversely with the
concentration of caffeine, caffeine citrate, and quinidine sulphate.
Sporulation failed to occur in all of the above solutions.

Table II shows the effect of different concentrations of strychnine
sulphate on the growth of Aspergillus niger at room temperature.
Column I shows the different concentrations of strychnine sulphate
used. The amount of growth obtained in each solution after 7-, 14-,
21-, 28-, and 35-day periods are given in columns II, III, IV, V, and
VI, respectively. Tables III, IV, and V show the effect of varying
concentrations of quinidine sulphate, caffeine citrate and caffeine,
respectively, on the growth of Aspergillus niger. These results were
obtained by the same method of experimentation and the data
treated the same as those in table II.

The amount of growth varied directly with the concentration
of strychnine and quinidine sulphate as shown in tables II and III.
The action of quinidine was different from that of strychnine in
that in the former there was considerable initial lag in the rate of
growth in solutions with concentrations above 0.15 percent. The
length of the initial lag, which varied directly with the alkaloid
concentration, reached a maximum value of 49 days in the highest
concentration used.

Table IV shows that caffeine citrate in concentrations of 0.05
percent or less may cause a slight increase in growth over the con-
trol solution, but in higher concentrations it seemed to have a rather
marked inhibiting effect. Caffeine produces a depressing action on
growth in concentrations of 0.05 to 1.00 percent, which increases in
direct proportion to the concentration as is shown in table V.

Table VI shows the effect of varying concentrations of different
alkaloids on the time interval in days required between inoculation
and the first appearance of growth and sporulation in cultures of
Aspergillus niger. Column I gives the concentration of alkaloid in
percent. Column II shows the number of days required for visible
growth to appear in solution with different concentrations of strych-
nine sulphate. Columns III, IV, and V show the same for solutions
of quinidine sulphate, caffeine citrate, and caffeine, respectively.
The time recjuired for sporulation in strychnine sulphate, quinidine
sulphate, caffeine citrate, and caffeine solutions of varying concen-
trations is shown in columns VI, VII, VIII, and IX, respectively.

The time required for the first appearance of growth and sporu-
lation in strychnine suli)hate solutions of all concentrations was the

96

The University Science Bulletin

same as that for the control sohition. In the sokitions of quinidine
sulphate, caffeine citrate and caffeine with concentrations above
0.10 percent, a longer time was required than in the control, the
length of time varying directly with the alkaloid concentration.
In the two highest concentrations of caffeine, sporulation was com-
pletely suppressed.

Figure I was taken from the results given in tables II, III, IV and
V and shows the maximum dry weight produced with each alkaloid
in each of the concentrations used. The alkaloid concentrations

eoo

CO

S 700

<r
a
0

-1 600

^/

^

^

y^

.3

^

^^'

^

"^

-^

\

i

^

0^

r
- 500

t-

X

w

5 400

0

N

sA

^^

■

\

5

--^

^^

^\

\

FIG.

0.1 o.t

16

a* 0.6 0.6 10 12. 14 1.6

ALKALOID COMCEMTRATIOM PERCE-MT
t-QUINlOIME- 2-STCYCHMiriEr 3-COMTCOL 4-CAr-fE-mE- CITRATE S-CAPffltlE-

: GRAPH SHOWIHG AMOUtlT OF- GROWTH IM DIFPtBtriT ALKALOID
SOLUTIOnS.

are given in percentages. The amounts of dry weight produced are
given in milligrams. This figure shows at a glance that strychnine
and quinidine sulphates increased, while caffeine citrate and caffeine
decreased dry-weight production. In the strychnine sulphate solu-
tions the amount of growth increased w4th increased concentration
of strychnine sulphate up to 1.80 percent, which was the highest
concentration used. In the quinidine sulphate solutions the amount
of growth likewise varied directly with the alkaloid concentration
up to 1 percent and then abruptly decreased. This abrupt decrease
in amount of growth may be traced to the exceedingly long initial
lag in the growth rate which is produced by high concentrations of
quinidine. From these results one may conclude that strychnine
and quinidine sulphates increase, while caffeine citrate and caffeine

Bates: Effects of Alkaloids on Growth

97

decrease the amount of growth. The action of strychnine and quini-
dine may be that of a stimiikis or nutrient or both. If these alka-
loids have a nutritive value, it may be either the nitrogen or carbon
or both that is assimilated.

Table I

Effect of different alkaloids on the growth of Rhizopus nigricans at room

temperature

I

II

Strychnine
sulphate.

III

Caffeine.

IV

Caffeine

citrate.

V

Qninidine

sulphate.

Alkaloid

concentration,

percent.

Growth
in grams
dry wt.

Growth
in grams
dry wt.

Growth
in grams
dry wt.

Growth

in grama

drj wt.

0.000

0.05

0.0112
0.0129
0.0152
0.0176
0.0194
0.0186
0.0256
0.0358

0.0112
0.0127
0.0106
0.0092
0.0086

0.0112
0.0104
0.0101
0 . 0096
0 . 0068
0.0021

0.0112
0.0095

0.10

0 . 0068

0.20

0 0059

0.30

0.40

0.50

0 . 75

T.ABLE II

Effect of strychnine sulphate on the growth of Aspergillus nigcr at room

temperature

I

Days

II

7

III
14

IV

21

V

28

VI

35

Alkaloid

concentration,

percent.

Growth

in grams

dry wt.

Growth

in grams

dry wt.

Growth

in grams

dry wt.

Growth

in grams

dry wt.

Growth

in grams

dry wt.

0.00

0 05

0.1342
0.16.34
0.1.596
0.1685
0.2220
0.1933
0.1844
0.2612
0.1517

0.3855
0 . 38.'^2
0.3922
0.3907
0.4738
0.4851
0 . 5046
0 . 4876
0.5484

0.5637
0.5603
0.5866
0.6043
0.6725
0.6616
0.7158
0.7410
0.7552

0.5710
0.5728
0.5933
0.5983
0.6111
0.6715
0 . 7060
0 . 7667
0.8017

0.5984
0 6162

0.15

0.30

0.50

0.75

1.01

1.40

1 . 80

0 . 5945
0.6178
0 . 6483
0.6810
0 . 7023
0.7890
0 7274

7—4141

98

The University Science Bulletin

Table III

Effect of varying concentrations of quinidine sulphate on the growth of
Aspergillus niger at room temperature

* First appearance of growth 49 days after inoculation.

I

Days

II

7

III

14

IV

21

V

28

VI

35

VII

42

Allvaloid

concentration,

percent.

Growth

in grams

dry wt.

Growth
in gram.s
dry wt.

Growth

in grams

dry wt.

Growth
in grams
dry wt.

Growth
in grams
dry wt.

Growth
in grams
dry wt.

0.00

0.1342
0.1686
0.1.536
0.1124
0.0674

0.3855
0.2320
0 . 2876
0.2747
0.3135
0.2004

0.5637
0.5677
0 . 5667
0 . 5704
0.5573
0.5624
0.1686

0.5710
0.6093
0.6044
0 . 6434
0.7011
0.5717
0.3953
0.1290

0.5984
0.6816
0.6717
0 . 6864
0.7334
0.7824
0.7005
0.1712

0 . 5883

0 05. . ...

0 15

0 30

0 . 50

0.75

0.7416

1 00

0.8616

1.25

0.6102

1 50*

Table IV

Effect of varying concentrations of caffeine citrate on the growth of
Aspergillus niger at room temperature

I

II

7

III

14

IV

21

V

28

VI
35

VII

42

Alkaloid

concentration,

percent.

Growth

in grams

dry wt.

Growth

in grams

dr.\- wt.

Growth
in grams
dry wt.

Growth

in grams

dry wt.

Growth

in grams

dry wt.

Growth

in grams

dry wt.

0.00

0.1.342
0.11.58
0.1210
0.1672
0 . 0920
0.0772
0.0.589

0.3855
0 . 3984
0.4126
0.4314
0.4100
0.3850
0.2667
0.1012
0.0364

0.5637
0.5810
0.5399
0 . 5248
0.4889
0.5011
0.5246
0.4356
0.3092

0..5710
0.5802
0 . 5290
0.4977
0.4754
0.5180
0.5321
0.4924
0.3991

0..5984
0 . 6056
0.5143
0.4621
0 . 4675
0.4928
0 . 52.57
0.4410
0.4205

0 . 5883

0 05

0.15

0.30

0.50

0 75

1.05

1.40

0 4985

1.80

0 4736

Bates : Effects of Alkaloids on Growth

99

Table V

Effect of varying concentrations of caffeine on the growth of Aspergillus niger

at room temperature

I

Days

II

7

III

14

IV

21

V

28

VI

35

VII

42

Alkaloid

concentration,

percent.

Growth

in grams

drj' wt.

Growth

in grams

dry wt.

Growth
in grams
dry wt.

Growth
in grams
dry wt.

Growth
in grams
dry wt.

Growth

in grams

dry wt.

0.00

0.05

0.1342
0.1396
0.1262
0.14.56
0.1708
0.0960
0,05.58
0.0346

0 . 3855
0.3092
0.3152
0.2791
0.2432
0 . 2308
0.1875
0.1258
0.1239

0.5637
0.5386
0..5330
0,3958
0.3843
0 . 3670
0,3210
0.2642
0 . 2280

0.5710
0.5662
0.5479
0.3963
0.4199
0.4198
0.3607
0 . 3422
0.2410

0.5984
0.5783
0.5240
0.3938
0 . 4046
0.3719
0,3464
0.3314
0.2744

0.5883

0.10

0.25

0.40

0.55

0.70

0 . 85

1.00

0.2917

Table VI

Time required in days for the first appearance of growth and sporulation of
Aspergillus niger at room temperature on different allcaloid solutions of
varying concentrations

Time required in daj
for first appearance of gr

•s
owth.

Time required in daj
for sporulation.

•s

I

Alkaloid

concentration,

percent.

II

SS

III

QS

IV
CO

V
C

VI

SS

VII
QS

VIII
CO

IX
C

0.000

0.100

0.625

1.175

1.750

1

1
1
2
5
180

1
1
2

5
12

1
1
2
2
5

3
3
3
3
3

3
3
5
8
183

3

3

11

15

34

3

3

25

SS — strj'chnme sulphate.
QS — quinidine sulphate.

CC — caifeine citrate.
C^caffeine.

100 The University Science Bulletin

NUTRITIVE VALUE OF STRYCHNINE
Material and Method

In the previous experiment it was found that strychnine and
quinidine sulphates accelerated and increased the growth of As-
pergillus niger. This experiment was undertaken to determine the
availability of strychnine-nitrogen and carbon.

Four different culture solutions were used. The different com-
pounds and amount of each used in the culture solutions are as
follows :

strychnine sulphate 15.00 grams

dextrose 20 . 00 grams

potassium nitrate 1 .48 grams

potassium dihydrogen phosphate 2.72 grams

magnesiimi sulphate 1 . 23 grams

distilled water to 1,000 c.c.

Culture solution No. 1 contained all of the above compounds.
The remaining solutions contained all of the above compounds with
the following exceptions: in solution No. 2, the strychnine sulphate
was omitted; in solution No. 3, the potassium nitrate was omitted;
and in solution No. 4, the dextrose was omitted.

The procedure used in sterilization, inoculating, and making dry-
weight determinations was the same as that used in the previous
experiments.

The amount of growth obtained in the different culture solutions
at the end of an 18-day growing period is given in Table VH. The
amount of growth represents the mycelium obtained from four cul-
ture flasks.

Table VII

Availability of strychnine-nitrogen and carbon

Solution
Number.

1
2
3
4

Carbon compounds
present.

Dextrose and strychnine.

Dextrose

Dextrose and strychnine.
Strychnine

Nitrogen compounds
present.

Potassium nitrate and strychnine.

Potassium nitrate

Strychnine

Potassium nitrate and strychnine.

Dry weight,
grams.

1.1415
1 .0846
0.0885
0.0009

The exceedingly small amount of growth obtained in culture solu-
tion No. 4 indicates that strychnine-carbon, when the only carbon
compound present, is unavailable. The small amount of growth ob-
tained in solution No. 3 indicates that strychnine-nitrogen can be
utilized to a slight extent in the absence of other nitrogen com-

Bates: Effects of Alkaloids on Growth 101

pounds. Good growth was obtained in solution No. 2 in which dex-
trose and potassium nitrate were used as sources of carbon and
nitrogen, and still better growth was obtained in solution No. 1,
in which strychnine was also present. From the results obtained in
this experiment one might attribute the increase in growth in the
presence of strychnine to the assimilation of strychnine-nitrogen
rather than to a stimulating action of the strychnine. This ques-
tion can be answered by studying the action of strychnine in cul-
ture solutions with varying ratios of dextrose and potassium nitrate.

To study this question a triangle experiment was set up using
fifteen different culture solutions. Each of the fifteen different
media contained 1.23 grams of magnesium sulphate and 2.72 grams
of potassium dihydrogen phosphate per liter. In addition the solu-
tion contained varying amounts of the following compounds: dex-
trose, an available form of carbon; potassium nitrate, an available
form of nitrogen ; and strychnine sulphate, which contains both car-
bon and nitrogen, the availability of which was to be studied. These
compounds varied in amount in such a way that each might be a
limiting factor in growth. The ratio of dextrose to potassium nitrate
varied in the different solutions from 1.25 to 80. Figure II shows
the dextrose, potassium nitrate, and strychnine sulphate content of
each of the fifteen different culture solutions in percentages. The
different culture solutions are referred to by numbers given below
the squares. 1,125 c.c. amounts of each of the fifteen different cul-
ture solutions were made up in 2-liter flasks and measured out in
75 c.c. portions into 125 c.c. Erylenmeyer flasks. After 30 minutes
sterilization in the Arnold steam sterilizer on three successive days
the ET-ion was adjusted to Ph 4.8.

The procedure for preparing the spore suspension and inoculating
was the same as that used in the previous experiment.

Dry-weight determinations were made with the procedure used in
the previous experiment at 11-, 28-, 41-, and 54-day intervals
following inoculation. The culture solution was removed from the
filter flasks and the latter rinsed with sufficient distilled water to
bring the solution up to its original volume. The culture solution
was then divided into aliquots which were used for H-ion and
quantitative sugar determinations, and a qualitative test for the
presence of nitrates.

Ph readings were made with a Sargent quinhydrone acid Ph meter
after heating the solution to expel any carbonic acid present. Quan-
titative sugar determinations were made by Benedict's method as
given by Mathews {SI ) . The presence of nitrates was indicated

102

The University Science Bulletin

Strychnine Sulphate, Dextrose, and Potassium Nitkate Content
OP THE 15 Culture Solutions in Percent.

/

SO. 125

A

/

d 2.00

\

. /

/

nO.20

\

\

15

S0.25

sO.25

/ d 1.50

dZ.OO \

/ n0.20

n 0. 15 \

\

13

M N

sO.75

sO.75

S 0.75

/ d 1.00

d 1.50

d 2.00 \

/ n O.ZO

hO.15

nO.lO \

1 10

11

12

\

/ S 1.25

Si.

25

S 1.25

S 1.25

\

/ d 0.50

d 1.

00

d 1.50

d 2.00

\

/ nO.ZO

nO.

15

r

»aio

nQ 05

\

/ 6 ?

8 9

\

/ S 1.75

S 1.75

S 1.75

S 1.75

S

1.75 \

/ d 0.25

d 0.50

d 1.00

d 1,50

d

2.00 \

/ n 0.20

nO.15

n 0.10

n 0.05

M

0.025 \

/ 1 2

3

4

5 \

Figure H

s, strychnine sulphate, d, dextrose, n, potassium nitrate

Each of the above solutions contains 0.123 percent of magnesium sulphate and 0.272
percent of potassium dihydrogen phosphate.

by adding a few drops of a solution of diphenylamine to a few c.c.
of the culture solution in a test tube and running chemically pure,
concentrated sulphuric acid down the side of the tube. A deep
blue ring at tlie juncture of the two liquids indicated the presence
of nitrate.

Results

Table VIII shows the amount of growth obtained with each of the
15 different culture solutions at the end of 11-, 28-, 41-, and 54-day
growing periods. Each weight listed represents the total mycelium
obtained from two culture flasks. The time intervals during which
sugar and nitrates were still present in the different culture solutions
are indicated by continuous and broken underlines, respectively.

Bates: Effects of Alkaloids on Growth 103

The results indicate that there was practically no further increase
in growth after the dextrose was exhausted in culture solutions Nos.
1, 2, 6, 7 and 10 (also 3 and 14, as indicated by their Ph values and
dextrose/KNOg ratios) in which the dextrose disappeared before
the nitrate. There was a gradual but slow increase in dry-weight
production after the nitrate was exhausted in cultures Nos. 4, 5, 9
and 12 (also No. 8, as indicated by its Ph value and dextrose/KNOj
ratio) in which the nitrate disappeared before the dextrose. This
would indicate that strychnine-nitrogen may be slowly assimilated
after the nitrate-nitrogen is exhausted, and does not preclude the
possibility of its being assimilated along with the nitrate. On the
other hand, the strychnine-carbon could only be assimilated along
with the dextrose, for there is no further increase in growth after
the sugar has been exhausted.

Table IX shows that there is a considerable increase in the acidity
of the solution when the nitrate is exhausted before the dextrose in
cultures Nos. 4, 5, 9 and 12 (also No. 8, as indicated by its Ph value
and dextrose/KNOg ratio). This may be due to the conversion of
the surplus sugar to organic acids (Klotz, 11^). The Ph value of the
solution remains higher when the dextrose is exhausted before the
nitrate in cultures Nos. 1, 2, 6, 7, and 10 (also Nos. 3 and 14, as in-
dicated by their Ph values and dextrose/KNOg ratios).

Table X shows, for each culture solution, the amount of sugar
consumed in grams per gram of fungus dry weight produced at the
end of 11-, 28-, 41-, 54-day growing periods. From the variation in
amount of sugar consumed per gram of fungus dry weight produced
in the different culture solutions it appears that either the carbon
of the strychnine is assimilated along with the dextrose to a greater
extent in some solutions than in others, or that the sugar is more
efficiently utilized in the production of fungus dry weight in some
cases than in others.

Table XI shows the amount of potassium nitrate consumed in the
production of one gram of fungus dry weight for several culture
solutions. The variation in amount would indicate that the nitrate
was reinforced to a greater extent with strychnine-nitrogen in some
solutions than in others Or that it was more efficiently utilized in
the production of dry weight in some cases than in others.

Figure 3 shows in graphical form: the percentages of potassium
nitrate, dextrose, and strychnine sulphate; the maximum amount of
fungus dry weight produced; and the average rate of growth in
milligrams per day for each of the fifteen different culture solutions.

104

The University Science Bulletin

-.045

.030

•.OIS

I 6 Z

5 0

10

6.6 7.5 10 10 (O IS 3

De-xtbose-Ano, Catio
7 13 II 3 15 14

eo
5

S IE 4 9

CULTuet SOLUT/OM NUWBE-K
S-STCYCHNIMt ti-Krt03 D-DtXTBOSE- G- GCOWIH R- BATE- OF- GROWTH

FIGURt 3: GRAPH COMPAPlhG AMOUNT OF GBOWTH AND COMPOSITIOri OP

nuTpiE-riT soLUTions

The numbers symbolizing the different culture solutions are the
same as those used in the previous figure and tables, and are given
in the lower row of figures below the graph. The ratios of dextrose
to potassium nitrate in the different culture solutions are given in
the upper row of figures below the graph.

The graph shows at a glance that there is but little if any correla-
tion between the amount of growth and the concentration of strych-
nine sulphate. The amount of growth shows a close correlation with
the dextrose concentration when the dextrose/KNOg ratio is below
13 and a close correlation with the KNO, concentration when the
dextrose/KNOg ratio is above 13. In the latter case the resulting
H-ion concentration of the culture solution is comparatively high as
compared with the former, table IX. This acidity may be due to
the conversion of the surplus sugar to organic acids (Klotz, 14)-

A more careful analysis of the graph shows that equal quantities
of dextrose in cultures Nos. 6 and 2 did not produce equal amounts
of fungus dry weight although dextrose was a limiting factor in
growth. This was also true of cultures Nos. 10, 7 and 3, which con-
tained equal amounts of dextrose. The greatest growth occurred in
the solution with a dextrose/KNOg ratio nearer the optimvun value
of 13.3. This is also true of potassium nitrate when it is a limiting
factor: cultures Nos. 9 and 4; and 8 and 12. In the latter two solu-
tions the smaller amount of growth was obtained in solution No. 8,

Bates: Effects of Alkaloids on Growth

105

in which the dextrose/KNOg ratio was closer to the optimum value
of 13.3. This may be due to two limiting factors in solution No. 8,
KNOg then dextrose, and only one limiting factor, KNO3, in solu-
tion No. 12. Analysis of solution No. 8 showed that the KNO3 and
dextrose disappeared at approximately the same time, table VIII.
That the KNO3 disappeared before the dextrose was shown by the
resulting H-ion concentration of the culture solution, table IX. So-
lution No. 12 had a greater amount of dextrose than solution No. 8.
After the disappearance of the KNO3, the fungus may have utilized
the stiychnine-nitrogen and continued to assimilate the surplus dex-
trose as was shown by a gradual increase in dry weight, table VIII
and figure III.

Table VIII

Amount of growth in grams of dry weight produced in each of the 15
different culture solutions

Days

11

28

41

54

Culture Number.

Weight
in

grams.

Weight
in

grams.

Weight

in
grams.

Weight

in
grams.

1

.1062

.1248

.1092

2

. 1602

.2278

.1866

.1919

3

.0962

..5370

.4786
.3556

.3928

t

. 1838

.3020

..3010

5

.1228

. 1432

. 1 508

.1648

6

.12^2

.2080

.1803

.1734

7

.1290

3656

. 5245

.4294

8

.2^20

.5018

.6318
.3095

. 5567

9

. 2956

.2952

.3057 •

10

.l.'S39

.3828

.4440

.3790

11

.317.=S

.4308

.4780

.4669

12 ^

.1Q18

6230

.6316

.6476

13

. 2.n66

.4400

.4972

. 5206

14

.'iO«?2

. 7938

.7207
.7146

.7423

15

.396«

.4688

.7345

'

106

The University Science Bulletin

Table IX

Hydrogen ion concentration of culture solutions taken at the end of the

growing period

Days

11

28

41

54

Culture Number.

Ph value.

Ph value.

Ph value.

Ph value.

1

2.95

2.90

2.87

2

2.75

2.35

2.30

2.70

3

2.60

2.20

2.00

*

2.40

4

2.50

1.90

*

5

2.45

1.90

*

*

6

2.50

2.37

2.28

2.30

7

2.64

2 20

2.28

2.38

8

2.45

1.90

*
*

*

9

2.20

*

*

10

2. 55

2.37

2.63

2.68

11

2.40

2 37

2.55

2.50

12

2.65

2 20

*

*

13

2.40

2.30

2.64

2.60

14

2 20

2.20

2 00
2.00

2.00

15

2.40

2.48

2.20

i

Ph too low to measure with potentiometer em^ployed.

Bates: Effects of Alkaloids on Growth

107

Table X
Dextrose consumed in grams per gram of fungus dry weight produced

Daj's

11

28

41

54

Culture Numpbr.

Grains

of

dextrose.

Grams

of

dextrose.

Grams

of

dextrose.

Grams

of

dextrose.

1

3.53

3.02

3.43

2

2.72

3.25

4.02

3.91

3

2.90

2.48

3.13
3.93

3.82

4

3.07

3.82

4.74

5

0.99

5.27

5.44

6.83

6

2.87

3.61

4.03

4.33

7

2.41

2.99

2.86

3.49

8

2.79

3.50

3.56
4.28

4.04

'

9

2.64

4.32

5.13

10

2.80

3.00

3.38

3.96

11

2.82

3.20

3.11

4.11

12

2.67

3.03

3.84

4.38

13

2.83

3.23

3.38

4.. 32

14

2.74

3.44

4.16
3.63

4.04

15

2.51

2.94

3.84

108

The University Science Bulletin

Table XI. — Potassium nitrate consumed, in grams, per gram of fungus

dry weight produced

Culture solu
number

tion

Grams of

KNO;,

Culture solution
number

Grams of
KNOa

3 .

4 .

5 .

279

210

227

9

12

14

242

231

283

8 .

237

QUANTITATIVE EXPERIMENT ON STRYCHNINE

ASSIMILATION

Material and Method

This experiment was undertaken to determine the amount of
strychnine removed by Aspergillus niger from culture solutions with
varying dextrose/potassium nitrate ratios.

The composition of the different culture solutions in percent
and their respective dextrose/potassium nitrate ratios are as follows:

Solution Number

Sugar/KNGj ratio.
Dextrose percent . .

KNO 3 percent

Strychnine percent.

MgSO^ percent

KH.jPO, percent . .

4

0.572
0.143
1.047
0.123
0.272

9
1.287
0.143
1.047
0.123
0.272

14

14
2.002
0.143
1.047
0.123
0.272

19

19
2.002
0.10.5
1.047
0.123
0.272

24

24
2.002
0.083
1.047
0.123
0.272

A control series with the above composition minus the strychnine
was run at the same time.

The culture solutions were made up in 1,800 c.c. amounts from
stock solutions.

All of the culture solutions showed a H-ion concentration of pH
4.33.

The culture solutions were measured out in 75 c.c. portions into
125 c.c. Erlenmeyer flasks and plugged with cotton. The solutions
were then sterilized and inoculated by the procedure used in the
previous experiment.

A few cultures were subjected, from time to time, to Fehling's
test for the presence of sugar and the test for the presence of nitrates
described in the previous experiment. These cultures were later
discarded. When the above tests indicated that either the dextrose
or nitrate had completely disappeared from the culture solutions,
the mycelium was removed, dried to constant weight, and weighed
as described in the previous experiment. Three cultures were used

I

Bates : Effects of Alkaloids on Growth

109

for each determination. The strychnine content of the culture
solutions was then determined by the following procedure: the
solution was made alkaline by the addition of ammonia, which pre-
cipitated the strychnine base in the form of crystals. This also pre-
cipitated a small amount of other substance from which the strych-
nine was separated by its solubility in chloroform. After stand-
ing overnight, the solutions were filtered through No. 589 filter
paper manufactured by Carl Schleicher and Schiill, the crystals
dried in a vacuum oven at 40 degrees centigrade, and weighed.
The strychnine portion of the precipitate was then removed by
running chloroform through the filter paper. The filter paper and
insoluble residue was dried to constant weight and reweighed. The
loss in weight represents the amount of strychnine in the culture
solution.

The accuracy of the above method for the quantitative deter-
mination of strychnine was found by adding equal amounts of
strychnine sulphate to 100 c.c. amounts of distilled water and to
100 c.c. amounts of different culture solutions in which the fungus
had been growing. The amounts of strychnine recovered from the
above solutions were as follows:

From distilled water
.7846 gram
.7809 gram
.7851 gram
.7827 gram

From culture solutions:
.7731 gram
.7825 gram
.7737 gram
.7845 gram

This shows a maximum experimental error of less than 2 percent.

Experimental Results

Qualitative tests for the presence of sugar and nitrate showed that
the dextrose disappeared before the nitrate in solutions Nos. 4 and
9 and that the nitrate disappeared before the dextrose in the remain-
der. The relative time required in days for the complete disappear-
ance of dextrose in solutions Nos. 4 and 9 and the disappearance
of nitrate in solutions Nos. 14, 19, and 24 are as follows:

Solution Number.

Strychnine
solution.

Control
solution.

4

16 days
18 days
16 days
12 days
12 days

22 days

23 days
22 days
18 days
16 flavs

9

14

19

24

110

The University Science Bulletin

The amounts of fungus dry weight produced and strychnine as-
similated in grams for each of the five different culture solutions
are as follows:

Solution Numbei

Growth in strychnine solution

Growth in control solution

Percent increase in strychnine solution
Strychnine assimilated

.3866

.2817
37.20
.0000

9

.6856
. 5573
23.00
.0000

14

.8822
.8011
10.10
.0360

19

.8649
.7403
16.80
.0036

24

.7095

.6477

9.50

.0000

The above data indicates a slight assimilation of strychnine in
solutions Nos. 14 and 19, while no change could be detected in the
strychnine content of the remaining solutions. The percentage in-
crease in growth appears to be greater in the strychnine solutions
with a low dextrose/KNOg ratio, which would indicate that the
increase in growth is not due to the assimilation of strychnine-
nitrogen. The results of these experiments indicate that strychnine
has but very little, if any, nutritive value, its action being purely
stimulative.

CONCLUSIONS

Strychnine sulphate acts to increase growth of Rhizopus nigricans
and of Aspergillus niger in Coons nutrient solution and quinidine
sulphate has a similar effect on the latter organism. Growth of
both fungi is decreased by caffeine citrate, and that of R. nigricans
by quinidine sulphate. These three chemicals also retard sporu-
lation by A. niger. Since A. niger makes no use of the carbon and
only slight use of the nitrogen contained in strychnine, increase
in growth of this fungus is believed to be due to a stimulatory action
resulting in efficient utilization of the carbon and nitrogen of the
nutrient solution.

LITERATURE CITED

(1) BoKORNY, Th. 1896. Vergleichenden Studien iiber die Giftwirkung ver-

schiedener chemischer Substanzen bei Algen und Infusorien. Pfliig.
Arch. Bd. 64: 302.

(2) BoucHARDAT. 1846. De I'influence du sol relativement a Taction des

poisons sur les plantes. Compt. Rend. Acad. Sci. Paris, 22: 674-675.

(3) CiAMiciAN, G., et Ravenna, C. 1912. Recherches sur la genese des

alcaloides dans les plantes. Ann. Chim. Phys., (8e ser.), 25: 404-
421.

(4) Cl.\utriau, G. 1894. Localisation ot signification des alcaloides dans

quelques graines. Ann. Soc. Beige de Microsc, 18: 35-54; 1906.
Recueil Inst. hot. Errera, 2: 265-280.

Bates: Effects of Alkaloids on Growth 111

(5) . 1900. Nature et signification des alcaloides vegetaiix. Ann. Soc.

roy. des Sc. med. et nat. de Bruxelles, 9: fasc. 2, 113 pages; 1902.
Recueil Inst. bot. Errera, 5: 1-87.

(6) CoMERE. 1910. Du role de I'alcaloide dans la nutrition des Algues. Bull.
Soc. Bot. de France, 57: 277.

(7) CoRNEViN, Ch. 1891. Action des poisons sur la germination des graines
des vegetaux dont ils proviennent. Compt. Rend. Acad. Sci. Paris.
113: 274-276.

(8) De Toni. 1905. Letter to Lutz. Bull. Soc. Bot. de France, 5: 201.

(9) De V.-\rigny. 1892. L'atropine est-elle un engrais vegetal? Rev. gen. de
Bot., 4: 407-420.

(10) Ehrlich, F. 1917. Ueber die vegetation von Hefen und Schimmelpilzen
auf heterocyklischen Stickstoffverbindungen und Alkaloiden. Bio-
chem. Zeitschr. Vol. 79: 152-161.

(11) GoppERT, Citation by Nobbe. 1874. Handbuch der Samenkunde, Berlin.

(12) GoRis, Albert. 1914. Localisation et role des alcaloides et des glucosides
chez les vegetaux, Paris. 2d edition.

(13) Hass and Hill. 1913. "An Introduction to the Chemistiy of Plant Prod-
ucts," 401 pp., Longmans, Green and Co.

(14) Klotz, Leo. J. 1923. Some aspects of nitrogen metabolism in fungi.
Annals of the Missouri Botanical Garden, Vol. 10 : 299.

(15) Lutz, M. L. 1898. Sur la nutrition azotee des plantes phanerogames a
I'aide des amines, des ,sels d'ammoniums composes et des alcaloides.
Compt. Rend. Acad. Sci. Paris, 126: 1227-1229.

(16) . 1899. Recherches sin- la nutrition des vegetaux a I'aide de .sub-
stances azotees de nature organique (amines, sels d'ammoniums
compo.ses et alcaloides), Paris, 1898 (these soutenue en 1899) ; Ann.
Sc. nat Bot., (8e ser.), 7: 1-103.

(17) . 1903. Sur le role des alcaloides envisages comme source d'azote

pour les vegetaux. Bull. Soc. Bot. de France, (4e ser.), 3: 118-128.

(18) , 1905. Sur I'emploi de substances organiques comme source d'azote

par les vegetaux vasculaires et cellulaires. (Resume), Bull. Soc.
Bot. de France, (4e ser.), 5: 194-202.

(19) M.ARC.^cci. 1887. L'azione degli alcaloidi nel regno vegetale ed animale.
Ann. d. Chim. ed Farm., Milano, 5: 3-7.

(20) M^RCET. F. 1825. De Taction des poisons sur le regne vegetal. Ann.
Chim. Phys. (2e ser.) 29: 200-224.

(21) M.ATHEWS. 1920. Physiological Chemistry. Third edition. William
Wood and Company, New York.

(22) MiRANDE. 1900. Recherches physiologiques et anatomiques .sur les Cus-
cutacees. These Doct. Sc, Paris.

(23) Mosso. U. 1894. Azioni di alcuni alcalloidi sul germogliamento del
semi e sul sucessivo sviluppo della pianta. Att. d. Soc. Ligustica
di scienze nat e geogr. Geneva, 5: 1-8.

(24) NoBECorRT. Pierre M. 1921. Action de quelques alcaloides sur le
Botrytis cinerea Pers. Compt. Rend. Acad. Sci. Paris, 172: 706-708.

(25) Otto. R. 1894. Welchen Einfluss haben Strychninesalzlosungen auf die
Entwickelung von Pflanzen in Sand und Humusboden? Naturw.
AVochensch., 9:625-626.

(26) Overton. 1897. Ueber die osmotische Eigenschaften der Zelle in ihrer
Bedeutung fiir die Toxikologie und Pharmakologie. Zeitschr. f.

Physikal Chem., 22: 189-209.

(27) Princeps, M. 1828. Note sur I'empoisonnement des vegetaux par les
substances veneneuses qu'ils fournissent eux-memes. Ann. Chim.
Phys., (2e ser.), 39: 95-97.

112 The University Science Bulletin

(28) Reveil, p. O. 1865. De Taction des poisons sur les plantes. These

Doct. Sc. Lyon. Paris. 97-122.

(29) Sawa, S. 1902. Are caffeine and antipyrin in high dihitions poisonous to

plants? Bull, of the College of Agric. Tokyo, 4: 411-412.

(30) Stracke, G. F. 1907. Onderzoekingen over de immuniteit van hoogere

planten voor haart eigen vergift. Proefschrift, Amsterdam; Archiv.
neerland. des Sc. exactes et naturelles, (2e ser.), 10: 8-61, 1905.

(31) Suzuki. 1901. On the localization of theine in the tea leaves. Bull, of

the College of Agric. Tokyo, 4: 297-298.

(32) TuNMANN. 1910. Ueber die Alkaloide in Strychnos nuxvomica L. wah-

rend der Keimung. Arch. d. Pharm., 284: 644-657.

(33) Weyl, Th. 1881. Ueber den Einfluss chemischer Agentien auf die As-

similationsgrosse grtiner Pflanzen. Sitz. Erlangen Phys. med.; 1882.
Analyse Centralbl. f. medizin. Wissensch., 20: 634-644.

(34) WiNTERSTEix and Trier. 1910. Die Alkaloide, Berlin.

(35) Wolf and Knop. 1865. Ueber die stickstoffhaltigen Nahrstoffe der Pfian-

zen. Die Landwirthsch. Versuchsstationen. 7: 463.

(36) Wood, Remington, and Sadtler. 1907. United States Dispensatory, nine-

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(37) Yasuda. 1901. On the effects of alkaloids upon some moulds. Bot.

Magaz. Tokyo, 15: 79-83.

(38) Young and Bennett. 1922. Growth of some parasitic fungi on synthetic

culture media. Am. Jour. Bot. Vol. 9. pp. 459-469.

This work was carried out in the Botany Department of the University of Kansas in
partial fulfillment of the requirements for the doctorate. The writer desires to make his
appreciative acknowledgment to Dr. W. H. Horr for his direction and encouragement during
the preparation of the work here presented.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV J June 1, 1938 [No. 3

The Flora of the Sand Hills of Harvey County, Kansas

JACOB HOMER DOELL,
Department of Botany, University of Kansas

Abstract: This study of the flora of the Harvey county, Kansas, sand
hills was made during the years 1932, 1933 and 1934. Special attention was
given to the factors which affected the plants, and the effect of the various
plants and other factors upon the topography of this area. Plants of the
various formations and the order in which they appear in these formations
are noted. This covers the various stages in the development of blowouts
and deposits, from their start until they reach their climax vegetation.

INTRODUCTION

IN a cursory survey of the plants of Harvey county, Kansas, the
vegetation of the sand-hill area presented such a striking differ-
ence in appearance from the surrounding territory that it was
deemed worth while to make a closer study of the causes for this
wide difference in the vegetation, so obvious even to the casual ob-
server. A search of the literature on this subject showed that the
sand-hill flora had not received the deserved attention in this state.
Other phases of plant life have attracted much wider attention than
this problem, which has been subjected to much intensive study in
other parts of the country. The study by Hitchcock (8), in his
Ecological Plant Geography of Kansas, is all that is directly related
to the problem.

The work by Cowles (5) on the Ecology of plants of the sand
dunes of Indiana and Michigan; the works of Pond and Clements
on the Phytogeography of Nebraska (11) and that by Rydberg
(13) on the vegetation of the sand-hill regions in Nebraska; and
finally, that by Gleason (7) on the vegetation of the inland sand
deposits of Illinois, were further inducements to undertake a more
intensive study of the ecological relationship of the plants of this
section. The material presented in this paper is from observations
made during the years of 1932, 1933, and 1934.

8—4141

(113)

114

The University Science Bulletin

McPhersonlfiVAJ.

Fig. 1. Taken from The State Geological Survey of Kansas, 1896.

SOIL

The soil was identified by The University of Kansas Geological
Survey of 1896 (9) as belonging to the Tertiary Eqims beds. The
sand is a surface residue due to erosion of this formation. There are
other and much more extensive sand-covered areas running west
and southwest across the state along the south side of the Arkansas
river.

PHYSIOGRAPHY

The topography of the region is extremely level, with no elevations
of any consideration, and gradually rises in the westerly direction,
starting with an elevation of 1,454 feet at Newton and rising to
1,535 feet at Hutchinson, a distance of about forty miles. The sec-
tion of most intensive study is about halfway between the two
points, near the town of Burrton and some 1,492 feet in elevation
above sea level.

Doell: Floka of the Sand Hills

115

CLIMATE

The climate is characterized by great extremes in wet and drought,
heat and cold; and, due to the level, unbroken surface, in preva-
lence of periodic storms, some of which are of considerable duration.

The average annual precipitation and the distribution of the
same by months as shown by the graph (fig. 2) is such that late
summer plants are discouraged during their blossoming and fruiting

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fit. Oa

r No.

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Fig. 2. Precipitation 1933,

1934,

Average precipitation 1897-1933, .

season. The graph also shows that the last two years of study have
been abnormally dry for this region. This may account for a pos-
sible reduction in the number of species found.

The wind. has an unbroken sweep, and blows from the south and
southwest throughout the spring and sometimes into early summer.

The extremes of heat and cold are very great and, when accom-
panied by strong, dry winds, make conditions very difficult for plant
life. Figure 3 shows the average monthly temperatures of Newton
for the last thirty-seven years and the deviations from this average
for the years 1933 and 1934.

]]6

The University Science Bulletin

SC"

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Fig. 3. Temperature 1933,

1934,

Average temperature 1897-1933,

EDAPHIC FACTORS

The sand deposit is naturally the great and determining factor in
the sand-hill flora and, therefore, was studied in some detail.

It is somewhat difficult to place the boundaries of this sand-hill
area at some places; at other places there is a sharp demarcation.

There has been considerable shifting of the dunes in the memory
of settlers in this region. Some dunes have moved north due to
prevailing southerly winds until they are now several hundred
yards away from their former position and in some places section
roads have had to detour these piles of sand.

The depth of this sand layer varies from the edge, where it may
be only a thin covering or intermingled with the subsoil, to the
highest dunes which are estimated to be from thirty-five to forty
feet high.

There are places at the bottom of blowouts, throughout the com-
plex, where the sand has been swept away to the subsoil.

The sand is characterized by a very marked uniformity in size
of particles. Table No. 1 shows the range of sizes of the sand
particles making up these dunes.

Doell: Flora of the Sand Hills

Table 1
Size of sand as determined by standard soil sieves

117

Number of screen.

20

40

60

80

100

.900

X
.900

.375

X
.375

.185

X
.185

.175

X
.175

.160

X
.160

Size of sand particles

.900
and
up

.375

to

.900

.185

to

.375

.175

to

.185

.160

to

.175

.160
and
less

Percent of soil retained by sieve

Trace

10

44.5

31.8

4.5

9.2

The No. 20 screen showed only a trace of particles too coarse to
pass through the .9 mm. x .9 mm. mesh and only 9.2 percent that
passed through the No. 100 screen.

Table 2

Comparative percolation rates of three soil types from this region

Time
percolation

starts.

Water percolating in 15-minute periods ( c.c).

Kind of Soil.

First
period.

Second
period.

Third
period.

Average.

Sand-hills sand

Garden loam

Sand-hills subsoil

3 minutes

30 minutes

4 hrs., 7.5 niin.

430
62.5
1.5

462
60

1.2

437
53.5
1.2

438
69
1.3

The percolation test of the soil shows the relative permeability of
this sand to water as compared with garden loam taken outside
the dune region and the subsoil of the dune region. The brass
cylinders used were 1 ^%g" in diameter and the column of sand was
8.5" high. This test shows the soil of the dunes to be so permeable
that no surface water drains off, even in the heaviest rains, but is
all taken up by the sand where it falls, and passes down through
the sand until it reaches the soil water level.

This table (No. 2) also' shows the rather impervious character
of the subsoil that underlies most of this sand-hill area. It is
less than 0.3 percent as pervious to water as the overlying sand.
This impervious property of the subsoil causes the formation of
ponds where the sand is blown down to the subsoil. These ponds are
quite numerous in wet seasons and some of them, in the deepest

118

The University Science Bulletin

blowouts, are so permanent that hydrophytic species are not un-
common in this otherwise typically arid area.

With the pervious condition of the sand and its low water-holding
capacity in mind, due to the size of the sand particles, and largely
to its lack of organic matter, the test for capillarity was applied
(table No. 3). This shows that the sand can readily supply its

Table 3

Test for capillary rise of water in the soils (in centimeters)

Kinds of soil.

Time Run.

Sand-hills
sand.

Garden loam
for comparison.

Sand-hills
subsoil.

5 minutes

6H cent.

9H cent.
IIM cent.
12 cent.
12H cent.
13J^ cent.
14J^ cent.
14H cent.
14% cent.
14% cent.
15 cent.

2 cent.

3M cent.

4}4 cent.

5 cent.

5% cent.

7% cent.
13K cent.
14% cent.
15K cent.
16 cent.
16M cent.

IK cent.
214 cent.
3H cent.
4J^ cent.

15 minutes

30 minutes

45 minutes . .

1 hour

2 hours

614 cent.

1 day

17H cent.

2 days

22% cent.

3 da vs

26 cent.

4 days

28 cent.

5 days

30 cent.

vegetation with sufficient water from the supply above the subsoil
only after the plants are well rooted.

The sand is so fine and has so little binding material mixed with
it, that it forms a loose mulch over the surface of the dunes when-
ever it dries. This helps to conserve the soil water against evapora-
tion, and keeps the sand moist to within six inches of the surface
even in the driest season, except in the places where dry sand is not
being deposited by the wind. This feature greatly reduces the
loss of water from the soil by evaporation.

HUMUS

The organic content of the sand, after thorough drying, averages
only .51 percent by weight. The method used for determination of
organic matter was that given by Braun-Blanquet (2) for soils
with no lime and little clay. A sample of the sand was dried in a
drying oven for three hours at 110° C, weighed and then heated to

Doell: Flora of the Sand Hills 119

white heat in a platinum crucible for one hour. The loss resulting
from igniting the sand was used as a measure of the amount of
organic matter present.

WATER-HOLDING CAPACITY OF SAND

The amount of water the sand is able to hold was determined by-
weighing and measuring the amount of laboratory dried sand re-
quired to fill a cylinder and then soaking the sand by immersing
the cylinder in water for an hour. After draining until no more
water would drip from the cylinder, the weight of the water held
by the sand equalled 27 percent of the weight of the dry sand. This
equals 38.5 percent of the volume of the sand. Sand dried to a
constant weight, in the laboratory atmosphere, still held 0.1 percent
water.

SOIL ACIDITY

The sand and the subsoil tested according to methods mentioned
by Braum-Blanquet (2), using the Baily (1) hydrogen electrode,
showed a slight acid reaction.

VEGETATION

The western part of Harvey county is in the region of change
from the bluestem or tall-grass prairie to the short-grass, or buffalo-
grass, prairie.

This change is not entirely abrupt and is best seen in the original,
unbroken prairie that has been fenced and pastured. Severe pas-
turing seems to be one of the causes to bring about the change from
bluestem to buffalo grass, for the prairie that was not broken and
has been used for hay meadows is still of the tall-grass type, while
the adjoining pastures are largely of the short-grass type except
where the soil is loose, due to its sandy character. Few of the main
short-grass species are found in loose sandy soils. The short-grass
pastures are characterized by Buchloe dactyloides (Nutt.) Engelm.,
Bouteloua oligostachya (Nutt.) Torr., and Bouteloua hirsuta Lag.
The first was never found in the sandy soil, and the latter two
species only where the soil is clayey enough so as to pack quite hard.
This alone could account for the absence of these grasses on the
dunes.

There is little, if any, of the original undisturbed prairie left in
this section of the state. Close pasturing has changed the vegetation
to such an extent that if left to itself without pasturing for awhile

120 The University Science Bulletin

it has little resemblance to that which has been preserved for prairie
hay meadows. In such places a large number of weeds now form
the dominant vegetation. The following are the most numerous
in such places: Allium mutabile Michx., Rumex crispus L., Amaran-
thus retrojiexus L., Baptisia bracteata Gray, Euphorbia marginata
Pursch., Callirhoe digitata Nutt., Opuntia Rafinesquii Engelm.,
Oenothera biennis L., Apocynum cannabinum L., Verbena bracteosa
Michx., V. stricta Vent., Solamim rostratum Dunal., Plantago major
L., Vernonia fasiculata Michx., Ambrosia psilostachyg, Gray, Xan-
thium commune Britton, Achillea millefolium L., Cirsium lanceola-
tum L. Hill, Helianthus annuus L., and Taraxacum officinalis Weber.

Even the unbroken hay meadows, through cropping, have changed
so that the tall prairie grasses are hardly dominant at certain times
of the year. Psoralea digita Nutt. (B&G), and Amorpha canescens
Pursch., often obscure the grasses completely, from a distant view.
Nevertheless this is the true prairie province with no natural timber
or brush, except along the watercourses, and, to use Cowles' term,
the tall grass association is the climax formation. Therefore we
shall consider any approach to this climax condition as an indica-
tion that the sand-hill vegetation is becoming stabilized.

A considerable portion of the sand-hill marginal land is under
cultivation. Here crops generally have to be planted late because
of the shifting of the sand by the spring winds. Drifting sand often
fills lister furrows and covers up corn in early July. These border
fields do produce excellent watermelons and sweet potatoes, both
being planted late in the season.

The region of most intensive study in this problem was seven-
teen miles west and about one and one half miles north of Newton,
Kan. A main road passing through the sand hills at this point
makes it quite accessible, and because of the shifting of the dunes,
a space has been left unfenced to the side of the road. In places
this is as much as a quarter of a mile wide. This area was not pas-
tured and the vegetation was as little disturbed as any that could be
found. This area contains dunes, level places, and blowouts in all
stages of their development. Most of the plant specimens studied
were taken from this area.

One of the striking features of this region is the number of species,
and the abundance of some of these species, that are seen on the
way to the sand hills, but are only rarely, or never, seen in the sand
hills.

The surrounding pastures are generally full of Vernonia fasiculata,
Euphorbia marginata, and Poa pratensis L., but in the sand hills

Doell: Flora of the Sand Hills 121

this species of Euphorbia was never observed, while the Vernonia and
Poa were each seen once. Cultivated fields in the surrounding
country are generally marked by Helianthus annuus L., but this
species was not found in the cultivated fields of the sand hills.
Instead we find Helianthus petiolaris Nutt.

The whole prairie region of this part of the state is marked by the
absence of trees, except along the watercourses, and those that
have been planted in other places. Shrubs are also lacking, except
those that have found refuge under trees or in places bordering
water bodies. The sand hills, in contrast with the surrounding
treeless plains, are dotted with Populus Sargentii Dode and Salix
nigra Marsh. An occasional Catalpa sp. and Morus ruba L. are
also found. Prunus angustifolis Marsh., the sand-hill plum, often
covers areas to the exclusion of other vegetation. This shrub some-
times covers as much as ten percent of a pasture. Cephalanthus
occidentalis L. is the shrub of next importance and may be found
as single shrubs, in clumps, or in thickets which may cover as much
as fifteen square rods. This plant forms the densest cover of
vegetation to be found in these dunes. Cephalanthus occidentalis is
described in Trees in Kaiisas as "a swamp-type species found grow-
ing only in swampy locations or along the margins of streams
where an abundance of soil moisture is available." In this area
it is often found perched on dunes which are twenty feet in height.
As a rule it is found along the low places and its presence on the
tops of the dunes can be explained only by the fact that its growth
rate is fast enough to keep its tops above the incoming sand. In
this way its roots are in a perpetual water supply while its tops
are exposed to very xerophytic conditions.

Salix tristis Ait., a dwarf willow, which does not exceed the me-
dium grasses in height, is quite common. An occasional unidentified
species of Rubus, Rosa setigera Michx., Amorpha fragrans Sweet,
and A. canescens complete the list of shrubs commonly found grow-
ing in these sand hills.

A considerable portion of this sand-hill area is practically bare
of vegetation because of the shifting sand which is blown back and
forth as the wind shifts from south to north and back to the south
again. Thus the plants which try to get established in the open
places usually get covered by the drifting sand, or have the sand
removed from around their roots. We may therefore assume that the
sand hills which are now covered by vegetation were once piles of
loose, drifting sand. This makes the successions of plants the
most interesting ecological problem of this area. To answer this,

122 The University Science Bulletin

with any degree of accuracy, one should have accurate data from
a considerable number of years. In a limited time the closest ap-
proach to this seems to be the study of the various dunes and
blowouts which represent the various stages in their development.
Blowouts are numerous throughout this area and may be studied
from their merest inception to their stabilized condition. They
occur throughout the sand hills wherever the sand is deep enough
so that the wind can undermine the roots of the existing vegetation.
A blowout may be started along a cow path or any other place
w^here the vegetation cover becomes broken enough for the wind
to start moving the sand. When the sand is removed from the
roots of the plants they die and topple over, leaving more sand
exposed to the action of the wind. The sand so removed is carried
until it is deposited around other plants which may be eventually

Fig. 4. Diagrammatic longitudinal section through a typical blowout; 1.
Original level of the sand. 2. Windward slope. 3. Basin. 4. Lee slope.
5. Deposit.

buried by it. When once started, a blowout usually keeps on grow-
ing until the leeward dune breaks the wind until it can no longer
carry sand fast enough to keep the plants out, or, until some plants
which can establish themselves in such conditions come in and stop
the movement of the sand.

The accompanying illustration, taken from Gleason (7), shows
well the condition of the ordinary blowout.

In this region the spring winds are more destructive than the
summer winds, for they are much stronger and the sand has the least
in the way of protective vegetation. With the coming of warmer
weather and more moisture, the sand which is not too actively
moving is soon covered wdth vegetation. As the strong winds of
spring and summer are from the south, most of the blowouts ex-
tend from the south to the north with the windward slope to the
south. Some blowouts get started in tlie winter, by the north winds,
and they head to the south. Plate XIII, figures 1, 2, and 3, are
different views of the lee slope of a deposit at the south end of

Doell: Flora of the Sand Hills 128

blowout. The sand here has been brought in from the north.
Figure 1 shows a portion of this slope which is covered by a thicket
of Cephalanthus about four feet in height. This was taken in
July, 1932. Figure 2, taken from a different angle, in March, 1933,
shows the same plants after they had been buried to within one foot
of their tops by the north winds of a snowless winter. Figure 3
shows the same spot from approximately the same angle as that from
which figure 2 was taken, in May, 1933. In that short time the
Cephalanthus had grown a foot in height and had the sand fairly
well protected from the action of the wind.

The average blowout has its windward slope to the south. This
slope is usually quite steep and has little vegetation. The plants
found here are usually those which have been undercut by the
winds and have slid down from the level sand above. A few summer
annuals may come in on this slope. There is, however, little chance
for seeds to become covered to a proper depth and get the right
moisture conditions to insure their germination and the establish-
ment of plants.

The bottom of an active blowout provides a more favorable
place for the starting of new plants, but here one finds mostly
late summer annuals. The seeds are carried in and covered by the
less violent summer winds, and, with moisture conditions here more
favorable the plants are able to get a start. Only the hardiest
herbs are able to withstand the constant contact with the continously
moving sand. ' The number of species found in such places are gen-
erally few, with Cenchrus pauciflorus Benth, Diodia teres Walt., and
Cristatella Jamesii T. & G. being the one most commonly found.
The last two are especially well fitted for such conditions. Diodia
is entirely covered by coarse hairs which are little injured by the
sand, while Cristatella is covered with sticky glandular hairs which
soon become covered wuth a protective layer of sand.

The sides of the trough are usually the steepest parts of the active
blowout. Here the wind-driven sand very rapidly cuts the banks
and removes the sand from under the plants' roots. The sides are
often undercut, as the vegetation which can succeed in the sand hills
usually has a very extensive root system, which penetrates the loose
sand to a great depth, thus holding the top soil until it is undercut.
Surface plants which slide down from the top supply practically all
of the vegetation found on these slopes. (Plate XIV, fig. 1.)

The lee slope of the blowout is less steep than either the wind-
ward slope or the sides of the trough and may be cut in the first
stages of the formation, but it soon becomes inactive. This slope

124 The University Science Bulletin

serves as a highway for the sand in its transportation from the
trough to the deposit. It, therefore, is a poor place for young
plants to get started and so remains bare until summer, when the
windy season is over. At this time the seeds that have been
covered to a favorable depth germinate and the slope becomes
sparsely covered by plants. A few perennial grasses may start
and live through the following seasons, but the bulk of the vegeta-
tion is composed of summer annuals.

There may be deep-rooted plants of the previous cover surface of
the lee slope, and there may also be plants on the face of the deposit
slope that have been partially buried each spring and that come
through the sand the following summer. In this way some slopes
have perennial plants as a residue from a previous occupation.
Plate XIV, fig. 2, taken July 4, shows an early summer condition,
while Plate XIV, fig. 3, shows the same place on September 16.
Aside from the few perennial grasses the plants are almost ex-
clusively Heliotro-pium convolvulaceum Nutt. This plant is a sum-
mer annual. The seed generally germinates in June, and the seed-
lings are about two to three inches high in early July. They
generally have to start under adverse conditions, for if a strong
wind carries the dry sand over the surface wdth any great velocity
the young plants are severely cut by it, and the intense light and
heat on this southern slope are severe tests for young seedlings.

Temperature readings on June 8, 1933, at 2:30 p. jn. showed as
follows: on the south slope the surface sand was 142° F.; at five
inches down it was 95° F.; and at ten inches down it was 86° F. At
the same time the north slope of the deposit was as follows: surface
sand, 126° F. ; at five inches down, 95° F. ; and at ten inches down,
86° F. At this time the Heliotropium seedlings were just getting a
start and were mostly not over an inch high, with their lower leaves
next to the hot sand; in fact, many of the leaves were in contact
with the sand.

No attempt was made to measure the intensity of the light, but
with the reflection from the light, buff-colored sand, it was very
high. Camera exposures had to be cut down to one half normal to
prevent overexposures.

The lee slope of the blowout and the deposit generally has a
greater variety of plants than shown in Plate XIV, fig. 3. Species
in the order of frequency could be arranged somewhat as follows:
Heliotropium convolvulaceum, Diodia teres, Cenchrus pauciflorus,
Cristatella Jamesii, Froelichia jioridana (Nutt.) Moq., Eragrostis

Doell: Flora of the Sand Hills 125

pectinacea (Michx.) Steud., Calamovilja gigantea (Nutt.) Scribn.
& Merr., Ambrosia philostachya.

The north side of the deposit at the end of a blowout that is
caused by a south wind is the region of greatest plant growth. Here
the vegetation is protected somewhat against the south wind and
the excessive heat and drought. The only adverse condition here is
that the deposit may receive so much sand in a given time that the
plants will be buried faster than they can grow. If a deposit starts
around Populus, Salix or Cephalanthus, a dune of considerable
height may be formed, as these plants can stand being covered up
from the bottom, and their growth keeps pace with that of the dune.
Plate XV, fig. 1, shows this condition in which cotton woods form the
main growth. A few annuals are present, but the perennial grasses
make an especially luxuriant growth at the foot of this slope.

If the deposit reaches a considerable height and the trough is
deepened in proportion, it reaches a stage where the blowout is
checked by its own development, because the wdnd is checked and
the sand cannot be carried over the deposit any longer. Thus the
blowout has reached its natural limits and so becomes inactive and
in the course of time is stabilized by the succession of plants that
normally bring about the perennial grass cover of such loose sand
areas.

This simple form of blowout is not the commonest form; for
blowouts are constantly reaching into each other and this compli-
cates their analysis.

There is a general tendency to form series of sand ridges from
north to south. Many of these are from fifty to two hundred yards
wide, and from a quarter of a mile to a mile in length. This is a
complex of blowouts which are very difficult to analyze, yet they do
not contain anything that is not analogous to a part of a simple
blowout.

RECOVERY

We assume that the tall-grass, or bluestem, prairie association is
the climax plant formation of all this region of the state of Kansas
outside the sand hills, and if any part would be left undisturbed
by cultivation, cropping or grazing it would finally revert to the
original condition and that, therefore, any near approach to this
condition could be considered as a stabilization in the sand-hill
vegetation.

The abundance of trees and shrubs in the sand hills is due pri-
marily to the fact that bare sand dunes serve, and have in the past
served, as barriers against prairie fires.

126 The University Science Bulletin

The recovery of the loose, bare-sand areas by the tall, perennial
grasses constitutes a series of changes and successions of plant
formations.

1. The Basin Association

The floor or basin of the blowout is generally destitute of plants
in a well-established stage of its formation, and generally remains
very sparsely settled until the blowout has reached its natural
limits. As stated above, there may be a few remnants of plants that
have been carried in from the surrounding banks or even some that
have weathered excavation, but generally it is practically bare of
plants. (Plate XV, fig. 3.)

Plate XVI, fig. 1, shows a stage in recovery in which Diodia teres,
Cristatella Jamesii, a few sedges, and some grasses, including Era-
grostis pectinaceae and Cenchrus paucijiorus, form a very sparse
covering.

In Plate XV, fig. 2, we have a stage in which young poplars grow-
ing from the roots of Populus Sargentii are the main plants. These
young shoots come from roots of small trees on the sides of the
blowouts and often run seventy-five feet or more just under the
surface of the sand. A root may give off many shoots as it crosses
the basin. It is interesting to note that Cowles (5) makes the
observation that this does not happen in the Lake Michigan region
with Populus monilijera Ait. and P. halsamijera Du Roi.

There are sometimes young poplars in these situations that come
up from seed; however, for this to happen the blowout must be
deep, and close to the clay subsoil, so that the basin will hold water
long enough for the roots of these seedlings to penetrate the soil
to perpetual moisture. This condition is not very common in these
sand hills.

If there is no great excess of moisture the basin may become well
wooded. (Plate XVI, fig. 2.) However, in a prolonged wet spell
the basin is apt to fill with water and drown out the young trees;
or the grasses and weeds become thick enough that if a fire sweeps
through the dry vegetation it will kill most of the young trees and
in that case the grasses that are not seriously hurt by fire become
the dominant cover. The roots of these grasses will sod the soil and
the dead leaves and stems add to the humus cover of the sand
until there is a tall-grass prairie.

2. The Lee Slope Association

The loose sand of the lee slope is a much more difficult place for
most plants to gain a foothold. The best sand binder of all plants

Doell: Flora of the Sand Hills 127

in these sand hills undoubtedly is Calamovilja gigantea. This grass
produces enormous root stocks that grow many feet in length in the
loose sand, and send up stems at intervals. These stems readily
grow from six to seven feet in height and a quarter of an inch or
more in diameter, wdth twelve to fifteen stem leaves, eighteen to
thirty inches long, besides having a bunch of basal leaves. The root
system is exceedingly well developed and reaches far into the wet
sand of the deeper strata so that the grass grows w^ell in the dryest,
hottest weather. The stem and leaves of this grass are so hard
that they stand the sand blasts, and will not decay in a year, thus
holding the sand in which they grow and constantly collect the
surface-blown sand. (Plate XVI, fig. 3.)

The success of Calamovilja gigantea in coping with this loose,
drifting sand depends largely upon the fast rate of growth of its
underground rootstocks, its ability to recover from being buried by
sand, it enormous root system, which supplies it with sufficient
moisture in the dryest seasons, and its copious growth of hardy
leaves and stem above the ground. This latter growth soon produces
a cover over the sand and provides conditions of moisture and humus
in which other plants can get a start and they in turn help hold
the sand from further blowing.

Calamovilja gigantea is confined to this loose-sand area entirely,
and as the interstitial plants become numerous and sand well settled
it invariably loses out.

Next to Calamovilja gigantea, Panicuni virgatum L. ranks as a
sand binder though it is not a pioneer in the loose sand. Once
started it is more efficient in sodding the sand than Calamovilfa,
for it produces a denser root system, and will stay past the loose-
sand stage and grow in well-sodded prairie. It produces a much
denser growth than Calamovilfa, though it does not reach the height
and coarseness of the former. (Plate XVII, figs. 1, 2.)

What can be called dune lee slope in the complex dunes composes
a large part of the loose-sand area; and the plants found there
depend largely upon the age or stability of the sand deposit. Be-
sides the species mentioned there are a large number of other plants
in this formation, of which the following species are the most
characteristic: Pasplum ciliatij olium Michx., Panicum virgatum.
P. Scribnerianum Nash., P. lanuginosum Ell., P. praecociwn Hitchc.
& Chase, Setaiia imberbis R. & S., Cyperus bushii Britton, C.
Schiveinzin Torr., C. jiliculmis Coult., Cristatella Jamesii, Cassia
chamaecrista L., Tephrosia virginiana (L) Pers., Lespedeza capitata
Michx., Lechae villosa Ell., L. tenujolia Michx., L. intermedia

128 The University Science Bulletin

Leggett., Physalis virginiana Mill., Plantago Purshii R. & S., P.
virginica L., Diodia teres, Ambrosia psilostachya, Xanthium com-
mune and Helianthus petiolaris Niitt.

3. The Deposit Association

In the simple blowout the deposit generally takes a fanlike form,
due partly to the less intensity of the wind here, and partly due
to the veering of the wind off the straight course. In the complex
dunes the deposit is a rather irregular loose sand area, receiving
fresh sand with every wind that carries sand. Even the side winds,
which may partly fill the trough again, help spread the deposit
sand over considerable area. Therefore the limits between deposit
and sodded areas are poorly marked in most places.

Aside from the regular plant formation characteristic of the de-
posits, this is the region of ruderal species. Here many vagrant
weed seeds, blown in from the outside, may get a start even if condi-
tions are not favorable for fruiting and establishment of the species.
However, many common weeds in the fields surrounding the sand
hills have never been observed even in this area. In most cases
the permanent grasses are encroaching from all sides and sodding
over the loose sand, and if the blowout loses its activity the whole
area will become covered with the vegetation of the settled sand-
hill prairie.

The most common weed species of the deposit are: Croton glandu-
losus L., Cyloloma atriplicifolia (Spreng.) Coult., Mollugo verticil-
lata L., Helianthus petiolaris, Froelichia gracilus Moq., F. floridana
Coult. & Nels., Oxybaphus hirsutis Sweet., Apocynum cannabinum
L., Verbena bracteosa, Solanuni rostratum, Diodia teres, Ambrosia
trifida L., A. psilostachya. Euphorbia geyeri Engelm., Sideranthus
annuus Rydb., Salsola Kali L., Cristatella jamesii, Lepidium apet-
elum Willd., L. ruderale L., Physalis virginiana, Chenopodiutn album
L., Amaranthus graecizans L., A. blitoides Wats., A. retrojiexus,
Lactica scariola L., and Solidago leptocephala T. & G.

Of the annual grasses on the deposit the following are the most
common: Digitaria sanguinalis (L.) Scop., Festuca octoflora Walt.,
Hordeum pusillium Nutt., Bromus racetnosus L., Cenchrus pauci-
fiorus, and Seteria imberbis. These annual grasses are followed by
the perennial grasses, which are usually of the following species:
Calamovilja gigantea, Eragrostis secundijiora Pres., Elymus virgini-
cus L., Paspalum dliatifolium, Panicum praecodu^, Chloiis ver'ticil-
lata Nutt., Triplasis purpurea (Walt.) Chapm., Aristida purpurascens f
Poir., Sporobolus canovirens Nash., Agrostis hyemalis (Walt.) B.
S. P.

Doell: Flora of the Sand Hills 129

4. The Prairie Association

The ultimate prairie association of the sand hills never reaches the
climax prairie formation of the surrounding country, due to the
characteristics of the sand that constitutes the sand hills. The
sand is too loose to form the short-grass prairie of many of the
surrounding pastures; while many of the grasses of the tall-grass
prairie have not been found in the sand hills. Among these may
be mentioned: Boutelous curtipendula (jVIichx.) Torr., Agropyron
S7nithii Rydb., A. repens (L.) Beauv., Schendonnardus paniculatus
(Nutt.) Trelease., Elymus virginicus L. and Tridens fiavus (L.)
Hitchc.

The low, wet places of the sand hills are characterized by the
following grasses: Tripsacum dactyloides L., Andropogon virginicus
L. (Plate XVII, fig. 3), Echinochloa crusgalli (L.) Beauv., Alope-
curus geniculatus L., Sporobidus crj/pfandriis I Torr. I Gray, Spar-
tinia viichauxiana Hitchc.

SAND BINDERS

The sand binders arc of prime importance here, as elsewhere, in
the stabilization of loose-sand drifts, for the duty of holding the
sand rests upon them until the subsequent species can get started
and cover the sand surface. The grasses readily hold the first place
in this important work, as is shown by the observations in our
locality and also in those of workers in other parts of the world.

Warming fi^l names Psamnia arcnario as the most important
sand binder in the coastal regions of Denmark. Secondary sand
binders, according to his observations in coastal regions, are three
woody plants: Hippophae rhamnoides, Salix repens and Empetrum
nigrum. In the inland dunes in Europe he names Elymus arenaricus
as the most important of sand binders.

Braun-Blancjuet (i) ranks two grasses, Agropyrum and Calama-
grostis, as the most important sand binders for inland Europe; and
Aristida pungous and Euphorbia gouyoniana as the chief sand bind-
ers in the North African deserts; while for the sea strand areas of
p]urope he places Elymus europaeus, Ammophila arenaria, and
Cyperus capitatus in the order of importance.

Cowles (5) places Ammophila ariouUnaceae first as dune former
and also as a sand binder; of less importance he classes Agropyrum
rlasystachum, Elymus canadensis and Calamograstis longifolia.

Hitchcock (9), in his article on "Reclaiming Sand Dunes," ranks
Ammophila arenaria as first in both Europe and America; and as a
9—4141

130 The University Science Bulletin

second he mentions Elynuis arenaria as a sand binder along the sea
coasts. For the interior dunes in Central Netherlands and Denmark
he places Calluna indgaris, a heath-forming shrub, as the chief plant
in artificial sand binding.

Pond and Clements (10) give Rcdfielda jlcxuosa as the chief plant
that finally controls the blowouts in the Nebraska sand hills by
binding the loose sand in those blowouts. It is often associated with
M iihlenbergia piingens as a secondary and aiding species.

Gleason (7) ranks Rhus canadensis as the most important sand
binder and dune former in the inland sand deposits of Illinois, and
Panicum virgatum as only slightly less efficient, and Tephrosia
virginiana and Ceanothus ovatus as secondary in this respect.

From this paper of Gleason's one may judge that he has to deal
with considerably less loose sand than is the case along the shore
of Lake Michigan; and probably not the severe drought and storm
that the Kansas sand hills are subjected to.

As mentioned before, Calamovilfa gigantia and Panicum virga-
tum, and especially the first-mentioned, are the first to become es-
tablished in our loose sand before any other plants can gain a foot-
hold. Along with the last-mentioned grasses there are the shrubs:
Cephalanthus, Prunus, Salix, and the tree Populus sargentii that
serve as dune formers even if they have little power of binding the
8and with tlicir roots. To these last should be added Yucca glauca
Nutt., which, due to its clusters of evergreen leaves, makes an
excellent windbreak (Plate XVHI, fig. 2, and Plate XVIII, fig. 1),
and due to its enormous storage root system (plate XVIII, fig. 3)
act to prevent the sand from blowing readily wherever it makes a
dense growth.

CONCLUSIONS

1. The sand hills of Harvey, Reno and Rice counties are similar
to other sand hills in Kansas, especially westward along the Arkan-
sas river. They are characterized as typically inland deposits with
a limited amount of sand. It is only prevailing winds from one
direction that cause movement of the whole complex in a general
direction.

2. The extremes in temperature, wind, moisture and light make
conditions difficult for plant growth.

3. The vegetation is characteristically xerophytic. The grasses
are the predominant plant cover both as to number of species and
as to stand.

4. Best sand binder for this area is Calamovilfa gigantia, witii

Doell: Flora of the Sand Hills

131

Panicum virgatum second in iniportance. Tlic latter, however, is
not able to establish itself in actively moving sand.

5. Trees in this area are able to survive because of the impervious
nature of the subsoil and the fire barriers formed by the bare sand.

6. Trees and shrubs which succeed here are able to propagate
themselves by underground parts. Seed propagation of woody
plants is relati^•cly unimportant.

7. Yucca glauca is im})ortant in holding the sand after it has been
stabilized.

8. The sand hills present a true transition between the tall- and
short-grass prairies, for some species of each formation invade the
sand hills while other species never appear there. Thus the prairie
climax of the sand hills is never that of the surrounding country.

ANNOTATED LIST OF SPECIES

Pteridophyta
Filicales

Marsileaceae

1. Mai'silea vestita Hook. & Grev.

Equise tales
Eqisetaceae

1. Equisetum laevigatuni A. Br.

Sperm ATOP HYT.\
Najadales
Alismaceae

1. Sagittaria latifolia Willd.

2. Sagittaria graminea Michx.

Graminales
Gramineae

1. Tripsacuni dactyloides L.

2. Andropogon scoparius Michx.

3. Andropogon virginicus L.

4. Andropogon furcatus Muhl.

.5. Sorghastrum nutans (L.) Nash.

6. Digitaria sanguinaHs (L.) Scop.

7. Leptolonia cognatum (Schultes) Chase.

8. Paspaluni eihatifolium Michx.

9. Pan';cum \irgatum L.

10. Panicum luiacluicae Aslie.

11. Panicum lanuginosum Ell.

12. Panicum praecocius Hitchc. & Chase.

13. Panicum Scribnerianum Nash.

14. Echinochlea crusgalli (L.) Beauv.

15. Setaria imberbis R. & S.

16. Cenchrus pauciflorus Benth.

17. Stipa spartea Trin.

18. Aristida intermedia Scribn. & Ball.

19. Aristida purpurascens Poir.

20. Alopecurus gcniculatus L.

21. Sporobolus clandestinus (Spreng.)

Hitchc.

22. Sporobolus asper (Michx.) Kunth

23. Sporobolus cryptandrus (Torr.) Gray.

24. Agrostis hyemalis (Walt.) B. S. P.

25. Calamovilfa gigantea (Nutt.) Scribn.

& Men.

26. Sphenopholis obtusata (Michx.) Scrilin.

27. Koeleria cristata (L.) Pers.

28. Spartina Michauxiana Hitchc.

29. Chloris verticillata Nutt.

30. Bouteloua oligostachya (Nutt.) Torr.

31. Bouteloua hirsuta Lag.

32. Triplasis purpurea (Walt.) Chapm.

33. Eragrostis megastachya (Kot'ler) Link.

34. Eragrostis trichodes (Nutt.) Nash.

35. Eragrostis pectinacea (Michx.) Steud.

36. Eragrostis secundiflora Presl.

37. Poa pratensis L.

38. Festuca octoflora Walt.

39. Bromus racemosus L.

40. Hordeum jubatum L.

41. Hordeum pusillum Nutt.

42. Elymus virginicus L.

43. Elymus robustus Scrilm. A- J. C. Sm.

44. Elymus striatus Willd.

Cyperaceae

1. Cyperus inflexus Muhl.

2. Cyperus Schweinitzii Torr.

3. Cyperus Bushii Britton.

4. Cyperus strigosus L.

5. Eleocharis palustris (L.) R. & S.

6. Eleocharis acicularis (L.) R. & S.

7. Fimbristylis autumnalis (L.) R. & S.

8. Scirpus validus \'ahl.

9. Scirpus lineatus Michx.

10. Hemicarpha micrantha (Vahl.) Britton

11. Carex pennsylvanica Lam.

132

The University Science Bulletin

Annotated List of

Xyridales

Commelinaceae

1. Tradescantia rtflexa Raf.

2. Commelina virginica L.

Liliales
Juncaceae

] . Juncus tenuis Willd.

Liliaceae

1. Allium niutabile Michx.

2. Yucca glauca Nutt.

Amaryllidaceae

1. Hypoxis hirsuta (L.) Coville.

Iridaceae
1. Sisyrinchium albiduni Raf.

Orchidales
Orchidaceae

1. Ibidiuiii ceinuuiii (I..1 House.

Salioales
Salicaceae

1. Salix nigra Marsh.

2. Salix tristis Ait.

3. Populus Sargentii Dode.

Urticales
Urticaceae
1. Morus rubra L.

Polygonales
Polygonaceae

1. Eriogonum annuum Nutt.

2. Rumex altissimus Wood.

3. Polygonum aviculare L.

4. Polygonvnn ramosissimimi Michx.
f). Polygonum Mulilenhergii (Meisn.)

Wats.

6. Polygonum pennsylvanicum L.

7. Polygonum Persicaria L.

8. Polygonum hydropiperoides Michx.

Chenopodiales
Chenopodiaceae

1. Cycloloma atriiilicifolium (Sprtng.)

Coult.

2. Chenopodium all>un L.

3. Corispermum hyssoiiifolium L.
■1. Salsola Kali L.

Amarauthaceae

1. Amaranthus retroflexus L.

2. Amaranthus graecizans I,.

3. Amaranthus blitoides Wats.

4. Froelichia floridana (Nutt.) Moq.

5. Froelichia gracilis Moq.

Species — Continued

Nyctaginaceae

1. Oxybaphus hirsutus (Pursh.) Sweet.

Aizoaceae
1. Mollugo verticillata L.

Caryophyllales
Portulacaceae

1. Talinum rugospermum Holzinger.

2. Portulaca oleracea L.

3. Portulaca pilosa L.

Ranimculales
Eammciilaceae

1. .\nemone caroUniana AValt.

2. Delphinium virescens Nutt.

Papaverales
Papaveraceae
1. Argemone intermedia Sweet.

Ciuci ferae

1. IjCpidium apetalum Willd.

2. Lepidiimi ruderale L.

3. Sisymbrium canescens Nutt.

4. Arabis virginica (L.) Trel.

Capparidaceae

1. Cristatella Jamesii T. & G.

Resales
Rosaceae

1. Rosa setigera Michx.

2. Rubus sp.

3. Prunus angustifolia Marsh.

Leguminosae

1. Schrankia uncinata Willd.

2. Cassia Chamaecrista L.

3. Baptisia bracteata (Muhl.) Ell.

4. Baptisia australis (L.) R. Br.

5. Crotalaria sagittalis L.
f>. Melilotus alba Desr.

7. Hosackia americana (Nutt.) Piper.

S. Psoralea tenuiflora Pursh.

9. Psoralea digitata Nutt.

10. Amorpha canescens Pursh.

n. Amorpha fragrans Sweet.

12. Petalostemum villosum Nutt.

13. Petalostemum candiduni Michx.

14. Tephrosia virginiana (L.) Pers.

15. Desmodium illinoense Gray.

16. Lespedesa capitata Michx.

17. Strophostyles pauciflora (Benth.) Wats

Geraniales
Linaceae

1. Linimi floridanuni (Planch.) Trel.

Doell: Flora of the Sand Hills

133

Oxalidaceae
Oxalis violacea L.
Oxalis stricta L.

Geianiaceae

Geraiiivini caiolinianuin L.

Zygophyllaceae

Tiiliuliis terrestris L.

Polygalaceae

Pulygala sanguinea L.
Polygala verticillata L.

Eiiphorbiaceae

Croton glandulosus L.
Euphorliia Geyeri Engelm.
Euphorltia inarginnta Puish.

Malvales
Malvaceae
Calliih(ie digitata Nutt.

Violales
Cistaceae

Crocanthemum majus (L.) Britt.
Lechea intermedia Leggett.
Lechea villosa Ell.
Lechea tenuifolia Michx.

Violaceae

^'iola Rafinesquii Greene.

Opuiitiales
Cactaceae

Opuntia Rafiiipsquii Engelm.

Myrtales
Lythraceae

Aninjannia roccinea Rottb.
Lythnim alatiim Piirsh.

Onasii'aceae

Lud\ igia alternifolia L.
Ludvigia palustris (L.) Ell.
Oenothera rhombipetala Nutt.
Oenothera hnmifusa Nutt.
Oenothera lariniata Hill.
Oenothera .'^errulate Nutt.

Gentianale.s
Apocynaceae

Apocynum cannahinum L.

A.^clepiadaceae

.■\srlepias tuberosa L.
Asclepias amplexicalis Sm.
Asclepias verticillata L.
Acerates angustifolia (Nutt.) Dec.

AxxoT.\TED List of Species — Continued

Polemoniales

Boraginaceae

1. Heliotropium convolvulaceum Nutt.

2. Lithospermum Gmelini (Michx.)

Hitohc.

3. Lithospermum angustifolium Michx.

Vei'benaceae

1. Verbena hastata L.

2. V'erbena stricta Vent.

3. Verbena bracteosa Michx.

Labiatae

1. Salvia azurea Lam.

2. Salvia lanceaefolia Poir.

3. Hedeonia hispida Pursh.

Solanaceae

1. Solanuni nigrum L.

2. Solanuni rostratum Dunal.

3. Physalis heterophylla Nees.

4. Physalis virginiana Mill.

5. Datura Tatula L.

Scrophulaiiaceae

1. Linaria canadensis (L.) Dumont.

2. Penstemon Buckleyi Pennell.

3. Veronica peregrina L.

4. Gerardia paupercula (Giay) Britton.

Acanthaceae
1. Ruellia ciliosa Pursh.

Plantaginales
Plantaginaceae

1. Plantago Purshii R. & S.

2. Plantago virginica L.

Riibiales
Rubiaceae

1. Galium Aparine L.

2. Diodia teres Walt.

3. Cephalanthus occidentalis L.

Canii>anulales
Cucvirbitaceae

1. Cucurbita foetidissima H. R. K.

Coinpositae

\'ernonia fasciculata Michx.
Liatris sciuarrosa Willd.
Liatris punctata Hook.
Liatris spicata (L) Willd
Liatris kansana (Britton) Rydb.
Sideranthus annuus Rydb.

7. Solidago nernoralis Ait.

8. Solidago leptocephala T. & G.

9. Aster multifiorus Ait.

134 The University Science Bulletin

Annotated List of Species — Concluded

10. Erigeron annuus (L.) Pers. 20. Helianthus subrhomboideus Rydh.

11. Erigeron canadensis L. 21. Coreopsis tinctoria Nutt.

12. Antennaria neodioica Greene 22. Bidens involucrata (Nutt.) Britten.

13. Gnaplialiuin polycephaluni Michx. 23. Gaillardia lutea Greene.

14. Ambrosia trifida L. 24. Achillea Millefolium L.

15. Ambrosia psilostachya Gray. 25. Artemisia ludoviciana Nutt.

16. Xanthiuni commune Britton. 26. Cirsium lanceolatum (L.) Hill.

17. Lepachys columnaris (Sims) T. & G. 27. Lactuca scariola L.

18. Helianthus petiolaris Nutt. 28. Pyrrhopappus grandiflorus Nutt.

19. Helianthus Maximiliani Schrad.

BIBLIOGRAPHY

(1) B.AiLEY, C. H. A Simple Hydrogen Electrode, Journal of American Chem.

Soc. Vol. 42, No. 1, 1920.

(2) Braun-Blanquet, J. Plant Sociology, McGraw-Hill Book Co., 1932.

(3) Clements. Frederick Edward. Research Methods in Ecology, Lincoln,

Neb., 1905.

(4) Call and Schafer. A Laboratory Manual of Agriculture. McMillan Co.,

1916.

(5) CowLES, H. C. The Ecological Relations of the Vegetation of the Sand

Dune of Lake Michigan. Bot. Gaz.. 27:95-321, 1899.

(6) . The Physiographic Ecology of Chicago. Bot. Gaz., 31:73-102, 1901.

(7) Gleasox, Henry Allan. The Vegetation of the Inland Sand Deposits of

Illinois. Illinois State Laboratory of Natural History, Vol. IX, Oct.
1910, Article III.

(S) Hitchcock, A. S. Ecological Plant Geography of Kansas. Trans. Acad.
Sci., St. Louis, Vol. VIII, 8:55, 1898.

(9) Methods of Controlling and Reclaiming Sand Dimes. U. S. De-
partment of Agriculture, Bureau of Plant Industry. Bulletin 57. 1904.

(10) PlA\yoRTH, Erasmus. The Uniyersitv Geological Suryey of Kansas. Vol.

II, 1897.

(11) Pond, R.. and Clements, F. E. The Phytogeography of Nebraska. 1.

General Suryey, Lincoln, Neb., 1898.

(12) RiBEL. D. Edward. Geobotanische Untersuchungs Methoden. Berlin,

1922.

(13) Rydberg, p. a. Flora of the Sand Hills of Nebraska. Contributed U. S.

Nat. Herb., 3:133, 1895.

(14) Steiger, T. L. Structure of Prairie Vegetation. Ecology, Vol. XI. No. 1,

1930.

(15) Warming, Eug. Oecology of Plants, Oxfoid Uniyersity Press, London,

1925.

(16) Trees in Kansas. Kansas State Board of Agriculture, Topeka, Kan., 1928.

136

The University Science Bulletin

PLATE XIII

Fig. 1. A lee slo}>e of a deposit with Ccphakuithus occideitialis about four
feet high. Taken July, 1932.

Fig. 2. The same as above, taken from a diffei-ent angle, after three feet of
sand has been deposited by the north winter winds. Taken March, 1933.

Fig. 3. The same as above, taken in Mav. 1£33.

Doell: Flora of the Sand Hills

137

PLATE XIII

t

*^ t^ T'^ •

i'

.MM

138 The University Science Bllletin

PLATE XIV

Fig. 1. A steep .side bunk of a blowout.

Fig. 2. The lee slope of a blowout, taken July 4.

Fig. 3. Another view of the samf> .^lope. taken Sept. 16.

Doell: Flora of the Sand Hills

139

PLATE XIV

# .

i

'■ . '■

s

,* -

■ d*

t -

-W

1

V

2

■>

140 The University Science Bulletin

PLATE XV

Fig. 1. A lee slojie of the deposit.

Fig. 2. A basin with Populus Sargcntii, starting from long lateral
roots which are just beneath the surface of the sand.

Fig. 3. A basin with very few plants.

Doell: Flora of the Sand Hills

141

PLATE XV

-r;5^

V

^

2

%

^^- *

I

142 The University Science Bulletin

PLATE XVI

Fig. 1. A basin becoming well covered by vegetation.

Fig. 2. A basin well wooded.

Fig. 3. Calamovilja gigantca as a sand binder.

Doell: Flora of the Sand Hills

143

PLATE XVI

«* ^->^Ww- mamm >

-. f^-^

■'ftfW^

^4}:-

• ;"" !,--'« '

iM

144 The University Science Bulletin

PLATE XVII

Fig. 1. Paniciim virgatuin developing a heavy surface cover.
Fig. 2. A Panicum virgatutn mat being undercut.
Fig. 3. Andropogon virginicui^ in a low spot.

Doell: Flora of the Sand Hills

145

PLATE XVII

10—4141

146 The University Science Bulletin

PLATE XVIII

Fig. 1. Yucca glauca stopping sand.

Fig. 2. The beginning of a deposit with Yucca glauca acting as a windbreak.
Yucca glauca roots in the background.

Fig. 3. Yucca glauca bein<:' undercut with the enormous storage roots ex-
posed.

Doell: Flora of the Sand Hills

147

PLATE XVIII

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXVJ . June 1, 1938 [No. 4

Studies in Anaphylaxis XVI: Physiological Studies of
Histamine and Peptone Reactions in the Cat

PAUL KABLER,

Department of Bacteriology, Univer.sity of Kansas, Lawrence

Abstract: The author reports the re.'^ults of his iihysiological studies of
histamine and peptone shock in cats. He obtained satisfactory results with
both sodium amytal and ether anesthesia. The typical reaction in the cat to
a single, rapid intravenous injection of eithei- histamine or peptone was a three-
phase drop in blood pressure. The ma.ximum reco\'ery time following histamine
injection was 26 minutes, while in no instance was there a complete recovery
of blood pressure following peptone shock. Histamine effects varied from a
slight decrea.*e to a definite acceleration of the heart rate, while peptone pro-
duced either a moderate or extreme slowing of the rate. Histamine caused a
fleeting increase in kiflney volume, while peptone produced a marked decrease.
Likewise histamine caused a definite increase in intestinal pressure, while in
peptone shock there was a loss of muscular rhythm and a relaxation of muscle
tone observed in most of the experiments. The intracystic pre.ssure was either
unaffected or increased in both histamine and peptone shock. The coagulation
time of the blood was unaffected by histamine and prolonged only slightly
by peptone. A definite effect of narcosis was observed following histamine
injection.

SECTION I— HISTAMINE

T^ALE and Laidlaw (1) in 1910 described the symptoms of his-
-■--' tamine shock of the intact animal of several species. They
found in the anesthetized cat that injection of 0.25 mgm. histamine
per kg. body weight produced a slowing of the heart, which did not
act directly on the heart muscle. The pulmonary pressure was in-
creased due to arteriole constriction, while the systemic blood j^res-
sure was decreased. The limb volume and intestinal loop volume
increased as the systemic blood pressure fell. The kidney and
spleen exhibited a marked decrease in volume. Excised gastric and
intestinal strips responded with contraction in the presence of high

(149)

150 The University Science Bulletin

dilutions of histamine. The body temperature was not materially
reduced nor was the blood coagulation time prolonged. Dale and
Richards (3) and others (3, 4, o, 6) have contributed to our knowl-
edge of histamine shock. Feldberg (7), Smith (8) and Bally (.9)
have conducted investigations of histamine shock in the rabbit. The
action of histamine as summed up by Wells ilO) is, "It causes
bronchial spasm in guinea pigs, obstruction to pulmonary circulation
in rabbits and a fall of blood pressure in dogs. It causes marked
local urticaria of the skin in humans, and it docs all these things in
extremely minute doses."

With this data in mind it seemed of value to verify the physio-
logical responses to histamine injection, and to add to the knowledge
of the responses to peptone injection in the cat, thus to compare
them with the physiological responses of feline anaphylaxis.

Kxperimental

This series included six normal, healthy cats, which varied in
weight from 2,200 to 4,000 grams. The average weight was 2,830
grams. Four of the animals were females and tw^o were males. The
various physiological experiments were performed simultaneously on
each of the cats except in the first few the technique of recording
changes in kidney volume was not perfected.

Anesthesia : The animals of this series were given intravenously
55 mgm. sodium amytal per kilogram, for three of which the anes-
thesia was incomplete, so the trachea was cannulated and an ether
bottle attached.

Injections: All injections were made by means of a cannula tied
into the right femoral vein. Each histamine injection was followed
by five c.c. of warm Ringer's to wash the cannula free of histamine
and insure us that the entire injection had reached the circulation.

The techniciue employed was essentially that described by
Bally i9).

Results

The responses of blood pressure, coagulation time and rectal tem-
perature recorded in this study confirm those reported by Dale and
Laidlaw. Twenty-fi\'e hundredths (0.25) mgm. of histamine, when
injected intravenously, produced an average drop in blood pressure of

Submitted to tl^e Department of Bacteriology and Farulty of the Graduate School in
partial fulfillment for the degree of Doctor of Philosophy.

Note. — Studies in Anaphylaxis XVI and XVII belong to a series of investigations of
anaphylaxis being directed or" conducted by N. P. Sherwood and O. O. Stoland of the Uni-
versity of Kansas.

Kabler: Studies in Anaphylaxis XVI 151

56.9 percent. Subsequent to 67 percent of the histamine injections
the blood pressure curve followed that described by Dale and Laid-
law as the "three-phase drop." The time required for the blood
pressure to recover was 7.5, 17, 20, 20, and 26 minutes, respectively,
in the five animals that survived the initial injection of histamine.
The recovery time progressively shortened with succeeding injections.
There was no apparent difference in the physiological responses when
under amytal fi'om those of ether anesthesia.

Heart rate: Five of the six cats survived the first injection of
histamine. The heart rate was unchanged in two, decreased by 15
beats in one, and increased by 5 and 77 beats, respectively, in two.

Kidney volume: In our experiments a fleeting but definite in-
crease in volume was observed. The increase was immediately re-
placed by a marked decrease in volume as the blood pressure fell.

Intestinal pressure: A definite increase of intestinal pressure fol-
low^ed every injection of histamine. The increase in pressure was
usually evident within 30 seconds following the start of injection.

Intraeystic pressure: Four cats receiving eight injections re-
sponded with an increase of intraeystic pressure. Two cats which
received four injections gave no response.

Death and narcosis: Our findings confirm those of other investi-
gators. In animals dead of histamine shock, post-mortem examina-
tion showed the large veins distended, the right heart engorged and
the left heart almost empty. In animals under light anesthesia, the
injection of histamine caused a loss of existing reflexes, such as the
corneal refiex, etc.

Discussion

In our experiments there was no appreciable difference in the
response of cats to histamine injection, under amytal, ether, or mixed
anesthesia. The blood pressure response followed very closely that
described by Dale and Laidlaw as the "three-phase drop." However,
not all animals produced this type of curve. Apparently cats are less
susceptible to the second injection of histamine than to the first, as
judged by the time necessary for the blood pressure to be restored.
Heart rates were reduced in one, unchanged in two, and increased in
two cats following histamine injections. They returned to normal
as the animal recovered from the shock.

Histamine injection in the carnivora did not affect the coagulation
time of the blood or the body temperature. Both the coagulation
time and the body temperature decreased progressively tliroughout
The duration of the experiment.

152 The University Science Bvlletin

A fleeting definite increase of kidney volume was observed, which
was immediately replaced by a profound decrease in volume. It
suggests that the preliminary action of histamine on the renal vessels
of the cat was a dilatation followed by tin active constriction.

From the intact intestinal loop records, it was shown that tlie
intestinal smooth muscle of the anesthetized cat responded in a
similar manner as the same when excised and tested in warm Tyrode
bath.

Since following sixty-seven percent of the histamine injections
there was a definite increase in the intracystic pressure, it suggests
that this response was due to the action of histamine. This is in
opposition to the view of Dale and Laidlaw, who thought histamine
had no direct effect on the bladder musculature.

Conclusions

We feel that the foregoing simultaneous physiological studies of
histamine injection in the cat bear out the following conclusions:

1. That sodium amytal anesthesia may be satisfactorily used
in laboratory studies.

2. The typical reaction of the cat to a single rapid injection of
histamine is a three-phase drop in blood pressure. There is an ab-
rupt drop, interrupted by partial or complete recovery, followed by
a slow, gradual fall to the minimum pressure.

3. The intravenous injection of histamine usually jiroduces either
a definite acceleration or no change in the heart rate. In one third
or one fourth of the cats it produced a moderate decrease in the
heart rate. The rate returns to normal as the blood pressure is
restored.

4. That the injection of histamine does not exhibit any notice-
able effect on the coagulation of the blood.

5. That it produces no evident change of the rectal temperature
in the cat.

6. That histamine injection produces a fleeting dilatation of the
renal blood vessels which is immediately followed by what appears
to be an active constriction with subsequent decrease of kidney
volume.

7. That histamine injection i)rotluces an active increase of in-
testinal pressure.

8. That in two thirds of the cases histamine induces a definite
increase of intracystic jiressure.

9. That cats dying in histamine shock display an arterial blood
pressure near zero. That the large veins are distended, the right
heart engorged and the left heart almost empty.

10. A definite effect of narcosis is observed following histamine
injection in a cat.

Kabler: Studies in Anaphylaxis XVI 153

SECTION II.— PEPTONE

Schmidt-Mulheim ill) in 1880 discovered that when i)eptone
was introduced intravenously into the systemic cii'cuhition of a dog,
it produced the remarkable results of delaying the coagulation time
of the blood drawn soon after the injection, and at the same time
caused a very profound fall of blood pressure. Polletzer {1:2) and
Thompson (13) have contributed to our knowledge of peptone shock
in the dog.

Likewise several authors il, 14, 15, 16, 17) have worked out
valuable information concerning peptone shock in the guinea pig
and rabbit.

Underbill compared the response of the cat to that of the dog as
follows: "In the cat the characteristic symptoms are evoked some-
what less readily, larger doses being necessary to produce comj'ja-
rable results."

Olivecrona {I4) found that dilution of from 1-250 to 1-750,000
of peptone readily increased the tone of excise intestinal strips of
the cat. Dilutions of peptone less than 1-200 produced a marked
relaxation and complete loss of rhythmatic contractions. Emery
and Griffith {6} found the effects of peptone in the cat to be a
decrease in liver volume and a fall of blood pressure.

EXPERIMENT.\L

This series was composed of twelve normal cats, ranging in weight
from 1,650 to 4,600 grams with an average weight of 3,090 grams.
Seven were males and five were females.

The twelve cats received twenty-two injections of peptone. Witte's
peptone was used throughout and the solution was made fresh for
each day's experiments. A boiled ten-percent solution of peptone
in i)hysiological saline was cooled and centrifuged to remove the
precipitate. The clear supernatent fluid was then used for injection.

Dosage: The dosage for the first ten animals was 0.25 grams of
peptone per kilogram body weight. Four tenths grams per kilogram
was the dosage for the last two cats of the series.

Injection: All injections were made intravenously by means of a
cannula tied into the right femoral vein. Each injection was washed
in with sufficient warm Ringer's to make the total volume injected
eriual ten cubic centimeters.

Anesthesia: Six of the cats were anesthetized by an intravenous
injection of 55 mgm. sodium amytal i^er kilogram. The remaining
six were anesthetized with ether.

154 The University Science Bulletin

Methods for determining the mean arterial pressure, heart rates,
coagulation time of the blood, rectal temperature, kidney volume,
intestinal pressure, and intracystic pressure were the same as those
described in the section of histamine.

Results

Blood pressure: A "three-phase drop" followed fifty-nine percent
of the intravenous peptone injections. The percentage drop and
the time required to reach the minimum value were greater following
the first injection than after the second. The average drop of blood
pressure following the first injection of pejitone was 44.9 percent.
The average time required for the blood pressure to reach its lowest
level after the first injection was two minutes, twenty seconds. In
no case was the recovery of blood pressure complete after the first
injection.

The average drop which followed the second injection of peptone
into a cat was fifteen percent, and the average time consumed in
reaching the low i)oint was forty-five seconds.

Heart rate: Following the first injection six cats showed a
moderate slowing of the heart. Of these animals four showed a
slowing of six to nine beats per minute, while in two the heart rate
dropped eighteen and twenty beats, respectively.

The heart rates of two animals were unaffected, and three exhibited
an increase of rate varying from two to eight beats per minute.

The second injection of peptone consistently produced no effect
on the rate of the heart.

Coagulation time: The blood of three cats following peptone in-
jection showed an increase of coagulation of from one minute, thirty
seconds to three minutes. The blood of three cats displayed a de-
crease of coagulation time following the first injection, but exhibited
an increase of one minute after the second injection. The blood of
six cats showed a progressive decrease of coagulability following
each injection.

Rectal temperature: In every animal there was a slow, ]irogressive
decrease of rectal temiierature throughout the duration of the ex-
periment.

Kidney vohnue: An analysis of the kidney volume records
showed that for ten cats injected with ]ieptone there was a marked
decrease in volume of the kidney after the first injection. Five of
the eight cats receiving two injections showed an increase of kidney
volume after the second iniection. Neitlier of the two animals

Kabler: Studies in Anaphylaxis XVI 155

which received a single fatal injection of ]ieptone showed any
chanp:c in the volume of the kidney.

Intestinal pressure: The initial reaction of the intestinal smooth
muscle of twelve, cats receiving injections of peptone, was a loss of
muscular rhythm accompanied by a slight or marked relaxation of
tone in seven, a slight but definite increase of intestinal pressure in
two and no change in three.

It has been shown, then, that following 58.3 percent of the initial
peptone injections in cats there was a slight or marked decrease in
the tone of the intestinal musculature.

Intracystic pressure: Three cats which received six injections re-
acted with an increase of intracystic pressure following each injec-
tion. The remaining nine cats of this series showed no change in
the tone of the cystic musculature following any injection.

Autopsy: Post-mortem examination of the cats dying in peptone
shock revealed the veins of the portal system and the interior vena
cava distended with blood. The right heart was engorged and the
left heart was almost em{)ty.

Discussion

Reports of investigators concerning the reactions of the cat to
peptone injections are few. No systematic or correlated simulta-
neous obsei'vations have been found describing the responses of the
various organs.

The results as recorded show that the animals react in somewhat
a variable manner. Initial intravenous injections of peptone into
the cat always jiroduce a dro]:) in blood pressure. The majority of
animals respond with the "three-phase drop" as described elsewhere;
however, a few show only a single-phase drop following peptone
injection. There is never a complete recovery of blood pressure
following the first injection. In contrast to the reported response
of the dog, our findings showed a decrease in the cat's heart rate
in one half of tlie animals and an increase in one fourth following
the first peptone injection; apparently there is also little tendency
for the normal heai't rate to be regained.

It has been shown that the dog is much less susceptible to the
action of the second or third injection of peptone than to the first.
Our experiments with the cat indicate that it likewise is refractory
to second injections of iieptone. The average blood-pressure loss
following the second injection Avas about one third that following
the first injection. The heart rate is consistently unchanged after
the second or third iniection.

156 The University Science Bulletin

Some dogs do not exhibit a loss of blood coagulability following
intravenous injection of peptone. We have shown that following
about thirty percent of the initial peptone injections in the cat the
coagulation time of the blood is increased.

In 83.3 percent of the cases the first injection of jieptone produces
a marked decrease in the kidney volume of the cat ; however, follow-
ing the second injection of peptone the kidney shows an increase in
volume. This might suggest that the initial peptone injection had
altered the reactivity or permeability of the renal blood vessels.

The response of the intestinal musculature to peptone in 58.8 per-
cent of the animals is a decrease in tone and a loss of rliythmic con-
tractions.

In seventy-five percent of the animals no response of the bladder
is recorded following peptone injection. Nevertheless, a small per-
centage of animals in this series shows a slight increase of both in-
testinal and intracystic pressure.

It is shown that the rectal temperature is unaffected by peptone
injections.

A perusal of the results shows no correlation in the response mani-
fest and the anesthetic employed. It appears that sodium amytal
may be used in the study of experimental peptone shock without
fear of depressing or masking some part of the reaction.

Conclusions

From the foregoing correlated physiological studies of peptone
shock in the cat we feel warranted in drawing the following con-
clusions:

1. That following an initial injection of peptone in the cat, there
is a characteristic drop of arterial blood pressure from which re-
covery is not complete in one hour, with an apparent lessening of
susceptil)ility to further injections.

2. That the heart rate is decreased in one half and increased in
one fourth of the animals following the initial injections of peptone.
It does not regain its normal value in the duration of our experi-
ments.

3. That following the second injection the blood-pressure drop
is relatively small and that the heart rate is unchanged.

4. That a decrease in the coagulability of the blood occurred in
over tliirty percent of the cats injected.

5. That the response of the intestinal smooth musculature of 58.3
percent of the animals to peptone injections is a decrease in tone and
loss of rh\thmic contractions.

Kabler: Studies in Anaphylaxis XVI 157

6. That following seventy-five percent of the peptone injections
there is no change in intracystie pressure.

7. That in 83.3 percent of the cats studied there is a consistent
decrease of kidney volume following the initial peptone injection,
and usually an increase of volume following the second injection.

8. That there is no correlation in changes of rectal temperature
with jK'ptone injections.

9. Thixt in fatal peptone shock there is engorgement of the vis-
ceral veins together with tlie large trunk veins of the body.

10. That sodium amytal anesthesia can be used in the study of
experimental peptone shock.

BIBLIOGRAPHY

1. Dale, H. H., and L,\idlaw, P. P. J. of Phy.siol. 1910, 41, 318.

2. Dale, H. H., and Rkhakds, A. N. J. of Physiol. 1918. 52, 110.

3. D.ALE, H. H., and Laidlaw, P. P. J. of Phy.siol. 1919, 52, 355.

4. Manwaring, W. H., Monaco, Marino, H. D. J. of Immunol. 1923. 8, 217.

5. Rich. A. R. J. of Expcr. Med.. 1921, 33, 287.

6. Emery and Griffith. J. of Phar. and Exper, Therap. 1921, 42, 233.

7. Fei.uberg, W. J. of Physiol. 1927. 63, 211.

8. Smith, M. J. of Immunol. 1920, 5, 239.

9. Bally, L. H. J. of Immunol. 1929, 17, 191.

10. Wells, H. G. Chemical Aspects of Immunity, 1925, pp. 209.

11. .Schmidt-Mulheim, quoted from W . H. Thomjison. J. of Phvsiol. 1896,

20, 455.

12. Polletzer, S. .1. of Physiol. 1886. 7, 283.

13. Thompson, W. H. J. of Physiol. 1896, 20, 455.

J. of Physiol. 1899, 24, 396.

J. of Physiol. 1899-1900. 25, 1.

14. (3livecrona, H. J. of Phar. and P]xper. Therap., 1921, 17, 162.

15. Bally, L. H. J. of Immunol., 1929, 17, 207.

16. Bakhr and Pick.. Arch. f. Exper. Path. u. Pharm.. 1913, 74, 41.

17. ScHiLTz, \V. H. Hyg. Lab. Bull. No. 80, 1912.

THE UNIVEESITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV I June 1, 1938 [No. 5

Studies in Anaphylaxis XVII: Physiological Studies of

the Hypersensitive Cat

PAUL KABLER and N. P. SHEIRWOOD,

Driiartinent of Bacteiiolog>-, University of Kansas, Lawrence

Ab.sthact : The authors review the literature and call attention to the fact
tliat in the few studies made by others on anaphylaxis on the cat complex
proteins capable of producing the Brodie reaction have been used. By emjiloy-
ing crystalline ejig albumin they were able to avoid the Brodie reaction. While
the.y were unable to produce acti\'e sensitization to this antigen in a series of
twenty cats they were successful in obtaining passive sensitization in six ( ut
of fifteen cats injected with high-titered antiserum obtained from rabbits.
Their results indicated that the failure to produce active .sensitization was
due to the poor antibody respon.se of the cat to antigenic stimulation. In
the passively sensitized cat the anaphylactic response was characterized by
a jirofound drop in artci'ial blood pi'essure, a marked reduction in the heart
rate, a pronounced decrease in kidnej- volume, a definite increase or no
change in the intestinal and intracystic pressure. The coagulation time of the
blood was unaffected. The uterine horns from .'sensitized cats gave typical
Dale reactions. The anajuiylactic response did not correlate perfectly with
either histamine or peptone responses, although there were resemblances to
both.

SECTION 1.— ACTIVE ANAPHYLAXIS

BRODIE (/) in 3000 described the exaggerated sensitivity of
normal cats to intravenous injections of foreign proteins. Ap-
parently this type of reaction was peculiar to the cat, since the thresh-
old of toxic reactions was much higher for the dog, rabbit, or guinea
pigs. All investigators (2, 3, 4, 5) who have reported studies of the
anaphylactic phenomena in cats have used as sensitizers complex
protein material such as horse serum, sheep serum, dog serum, or
fresh egg white. These substances, even in small amounts, produce
a decided drop of arterial blood pressure when injected intravenously
in the normal cat.

(159)

160 The University Science Bulletin

Nevertheless, Manwaring (2) and Edmunds (4) have described a
slow drop in blood pressure in sensitized cats following the injection
of the shock dose, and thought is was characteristic of true anaphy-
laxis.

The ambiguous reports suggested the question as to the pos-
sibility of actively sensitizing cats with a chemically pure antigen
such as crystalline egg albumin. By the use of this antigen it was
hoped to eliminate the "Brodie Reaction" and to secure simultaneous
physiological responses of the various organs.

The question also arose as to the possibility of passively sensitizing
cats to crystalline egg albumin.

In all experiments the results were obtained under as nearly identi-
cal conditions as experimentation would ]iermit. The methods em-
ployed were the same as those reported in the preceding paper {6).

Experimental

This series was composed of twenty cats, all of which were grown
except one that was used for smooth muscle experiments alone. This
was a young female weighing 1,200 grams. The adult cats varied in
weight from 1,800 to 8,520 grams. There Avcre sixteen females and
four males included in the adult series.

Sensitization: The sensitizing antigen was a solution of four-jjer-
cent crystalline egg albumin (7) in distilled water. The sensitizing
dose was 0.5 c.c. of four percent albumin solution per Kg. of body
weight. Several methods of injection were utilized in attempting to
sensitize the animals of this series. The methods used were:

1. One subcutaneous injection followed at three-day intervals by
two intravenous injections.

2. Three intravenous injections given two days apart.

3. Eight to thirteen subcutaneous injections given in three periods
of three to five injections on consecutive days witli three days rest
between jieriods.

4. Three subcutaneous injections on consecutive days, followed by
an intravenous injection the fourth day, allow three days to elapse
and repeat.

5. Five to seven subcutaneous injections given at two-day inter-
vals and followed by two intraperitoneal injections of the same
spacing.

Incubation period: Periods varying from 8 to 74 days were in-

NoTB. — Studies in Anaphylaxis X\'I and X\'1I In-long to a series of investigations^ of
anaphylaxis being directed or romluftcd l.y N. P. Shi-rwond and O. O. Stoland of the Uni-
versity of Kansas.

Kabler and Sherwood: Studies in Anaphylaxis 161

vestigated. Most of the animals were tested 18 to 26 days after the
last injection.

Anesthesia: For each animal ether was administered from a cone
until complete surgical anesthesia was established, after which the
trachea was cannulated and an ether bottle attached.

Shock dose: The shock dose was 2 c.c. of four percent crystalline
egg albumin solution per kg. of body weight. It was injected by
means of a cannula tied in the right femoral vein. Sufficient warm
Ringer's solution was injected to make sure that all the albumin
had entered the circulation.

Precipitin titration: Samples of blood were drawn immediately
before and five minutes after the injection of the shock dose. Their
precipitin content was determines 1 by use of the ring test and over
night settling methods.

Resilts

Th.e blood pressure showed invariably an injection rise of from
four to twenty millimeters of mercury, which was maintained for
from 30 seconds to ten minutes. This was essentially the same re-
action as of a normal cat when injected with an identical dose.

The lieart rate usually exhibited slight slowing. The maximum
decrease of rate was ten beats per minute. The coagulation time
of the blood and the rectal temperature progressively decreased
throughout the duration of the experiments.

The intestinal and intracystic pressures were consistently without
change.

The kidney volume usually showed no change; however, a very
slight increase sometimes accompanied a pronounced injection rise
of blood pressure. A decrease in \-oIume never followed the injection
of the shock dose.

The intestinal strips from each of seven cats, and uterine horns
from six were removed and suspended in Ringer's solution kept at
37°C. under light tension until the ])hysiological experiments were
begun. At that time they were tested by the method of Schultz and
Dale for specific contraction with the homologous antigen, crystal-
line egg albumin. All the intestinal strips as well as the uterine
horns failed to respond whCn tested by this method.

Cluinea pigs were injected with the sera of six supposedly sensi-
tized cats in an effort to passively sensitize the pigs to crystalline egg
albumin. The cat sera were injected intraperitoneally in five cubic
centimeter amounts into 40()-gram normal-health guinea pigs.

11—4141

162 The Uxiversity Science Bulletin

After twenty-four to seventy-two hours incubation, the guinea
pigs were injected intracardially with one cubic centimeter of four
percent crystalline egg albumin and were observed for clinical symp-
toms of anaphylaxis. None of the guinea pigs responded with
anaphylactic symptoms.

The sera of twenty-one cats were tested for the presence of pre-
cipitins for crystalline egg albumin and all found negative.

Discussion

It is quite evident from the preceding results that no cat in this
series displayed any of the recognized symptoms of anaphylactic
shock. The possibility that this lack of sensitization was due to
nonantigenicity of the crystalline egg albumin had to be ruled out.
This was accomplished by at least three biological methods:

(a) Guinea pigs could be sensitized by a single subcutaneous in-
jection of one cubic centimeter of four percent crystalline egg al-
bumin. An identical dose when injected intracardially twelve days
later produced fatal clinical anapliylaxis.

( b ) Typical specific contractions of the excised uterine horns
from sensitized virgin guinea pigs were demonstrable by Dale's
technifiue.

ic) Multiple intravenous injections of crystalline egg albumin in
rabbits stimulated the {production of easily demonstrable jirecipitins
for that antigen. This method will be described in detail in the fol-
lowing section.

These findings seemed to prove the antigenicity of the crystalline
egg albumin. If this were true, the ciuestion arose as to why the
cats were not sensitized by its injection. It has previously been
pointed out that apparently the cat produces no precipitating or
passively sensitizing antibodies to crystalline egg albumin. Is there
likewise no hemolysins nor bacterial agglutinins produced in re-
sponse to introduction of their respective antigens? AMth the hope
of ascertaining the answer to this question, a series of three cats
Avas given five intraperitoneal injections of ten i^ercent sheep cells
at three-day intervals, and ant)tlier series of three cats was given
intraperitoneal injections of a formalized suspension of E. typhi
(Rawlin's strain) .

It was found for the first series that all the cat sera preceding
injection contained hemolysins for sheep cells when undiluted, but
not in a dilution of one to ten. After the round of injections there
was complete hemolysis of the sheep cells in a dilution of one to
ten, but only a trace in a dilution of one to fifty.

Kabler and Sherwood: Studies in Anaphylaxis 163

None of the animals of the second series sliowed any agglutinins
for E. typhi before immunization. After five injections the sus-
pension of E. typhi was agglutinated by 1-200. 1-100, and 1-32 dilu-
tions of the respective sera.

Thus it is shown that at best the cat produced antibodies in very
small quantities. It was thought that this might be explained by
an inadequate absorptive mechanism, or by a complete inability to
absorb foreign substances from the peritoneal cavity. However,
when trypan blue was injected intraperitoneally, it appeared in the
mucous membranes and skin as early as five hours after injection.

Three cats were then injected intraperitoneally with five cubic
centimeters of ten percent suspension of chicken cells. One of the
cats was killed after the elapse of six, twelve, and twenty-four hours,
the spleen and hvvv removed, embedded, sectioned, and stained.
Definite phagocytosis of the chicken cells was observed by the fixed
tissue cells of the spleen and liver from the cats killed at 12 and 24
hours. It a]ipears, then, that the cat has an adequate absorption as
well as phagocytic mechanism, although it does not respond with
active generation of demonstrable antibodies to the introduction of
foreign jM-oteins.

The apparent inability of the cat to generate antibodies may ac-
count for the entirely negative series just reported. It seems very
reasonable that if hemolysins, bacterial agglutinins, and precipitins
are not generated in demonstrable quantities, that likewise the
sensitizing antibody of anaphylaxis is also absent. This assumption
is certainly borne out by the preceding physiological studies of ana-
phylaxis.

SECTION 2.— PASSIVE SENSITIZATION

It has been pointed out that the literature concerning active
anai)hylaxis of the cat is very meager, thus it is not surprising that
the passive phenomenon is apparently omitted. We have offered an
explanation as to why the cat does not readily yield itself to active
sensitization. Howe^■er, it was shown that foreign substances are
quickly absorbed from the body cavities of this animal. So the
question remains as to the possibility of passively transferring the
anaphylactic sensitizer to the cat.

Experimental

This series was composed of fifteen adult, healthy cats, ten of
which were females and the remaining five were males. The animals
ranged in weight from 1,800 to 3,050 grams.

1

164 The University Science Bulletin

In each experiment the various physiological responses were re-
corded simultaneously with a few exceptions which will be noted
with their resjiective analysis. For each animal the anesthesia and
the methods of recording the various physiological responses were
the same as those described in the section on active sensitization.

Sensitization: High-titered anticrystalline egg albumin sera were
produced by repeated injections of four percent crystalline egg al-
bumin solution into the marginal ear vein of rabbits. Seven sera
were prepared which showed a precipitin titer of from 1-17,500 to
1-27,500 and which were capable of passively sensitizing 400-gram
guinea pigs when injected intraperitoneally in two cubic centimeter
amounts. An analogous dose, or five cubic centimeters per kg.
body weight, was chosen as the sensitizing dose for cats and was in-
jected intraperitoneally.

Incubation period: The incubation period varied from twenty-
four to seventy-two hours.

Shock dose: The shock dose, consisting of two cubic centimeters
of four-percent crystalline egg albumin solution per kg. of body
weight, was injected through a cannula into the right femoral ^-ein.
The cannula was then washed clean of antigen with sufficient warm
Ringer's solution to bring the total volume injected to ten cubic
centimeters.

Results

Blood pressi(.re: Six cats showed a profound drop of arterial blood
pressure following injection of the shock dose. Three of the cats
responded with a "three-phase" drop as previously described, and
three with a single-phase drop. The droj) (;f blood pressure was
initiated from thirty seconds to one minute and forty seconds after
the injection of the shock do'se was started. The average droi^ of
blood pressure was 42.7 percent. The low point of the curve was
reached on an average of two minutes, thirty seconds after injection
of the shock dose. The recovery time in the six cats was 4. 7, 8,
8, 20, and 28 minutes, respectively. There was no correlation
observed between the percentage drop in blood jn'essure and recovery
time, both the seven-minute and twenty-eight-minutc recovery
times being associated with fifty-percent drop in blood pressure,
while the four-minute and twenty-minute recovery times were as-
sociated with thirty-f()ur-i)ercent drops in blood pressure.

The second injection,* consisting of two cubic centimeters of four

* This is the term used to denote the injection of the specific protein to test for de-
sensitization of the animal.

Kabler and Sherwood: Studies in Anaphylaxis 165

percent crystalline egg albumin jicr kg. body weight, invai'iably
])roduced only a transient injection rise of blood pressvire. The
absence cf a bluud-pressure drop following the second injection
would suggest complete desensitization of the animal by the shock
dose of specific antigen.

Heart rate: Of the six positive animals, all but one exhibited a
decrease in heart rate during the period of lowered blood pressure.
In three of the animals the reduction of rate was marked, amount-
ing to 26, 81 and 91 beats, respectively; two showed a reduction
of 5 and 6 beats, res]-)cctively, while in one animal there was no
change of rate.

Coagulation time and rectal temperature: In no instance was
there a prolongation of clotting time associated with the specific
drop in blood pressure. The value of both the coagulation time
of the blood and the rectal temperature i)rogressively decreased
throughout the duration of the experiments.

Kidney voluvie: In every case there was a pronounced decrease
in the renal A'olume as the blood pressure fell. Usually the volume
of the kidney remained greatly diminished long after the blood
pressure had started recovery. In no case was there a decrease in
kidney volume following the second injection of albumin.

Intestinal pressure: Of the six cats responding with a drop of
arterial blood pressure, four exhibited a definite increase of intestinal
pressure following introduction of the shock dose. There was no
change of pressure following any of the second injections.

Intracystic pressure: Two animals showed a definite increase of
intracystic pressure after injection of the shock dose. There was
never any increase of intracystic ]n'essui'e following the second
injection.

Precipitin content and passively sensitizinej power for guinea pigs:
The sera of fifteen cats were titrated for precipitin content. Samples
drawn immediately before injecting the shock dose and five minutes
after injection were tested. Both the Ring Test and overnight
settling methods were employed. All the sera were consistently
negative, even when tested with undiluted antigen.

The passively sensitizing power of the cat sera for guinea pigs was
tested as has been described. The sera of three positive reacting cats
and one negative reacting cat were employed. A twenty-four hour
incubation period was allowed to elapse before injecting the shock
dose, which consisted of one cubic centimeter of four-percent albumin
solution. None of the guinea pigs displayed anaphylactic symptoms
following intracardial injection of the shock dose.

KJG The University Science Bulletin

Excised intestinal strips and uterine horns: Two intestinal strips
from each of seven negative cats and from two positive cats were
tested for specific contraction by the Schidtz-Dale technique. None
of the strips from the negative animals responded with a specific
contraction, while the strips from both positive cats reacted with a
marked contraction when one cubic centimeter of two-percent egg
albumin was added to the twenty cubic centimeters Tyrode's solu-
tion bath.

After renewing the bath with fresh Tyrode's, a second addition of
an identical dose of albumin caused no contraction. A subsequent
addition of barium chloride produced the characteristic shortening.

The uterine horns from three negative cats and from two positive
cats were tested by the same method. The horns from one positive
cat were hyperirritablc, relaxation was unobtainable and a satisfac-
tory test was impossible. The horns from the other positive cat re-
sponded witli a maximum contraction when 0.8 c. c. of two-percent
albumin was added to the 20 c.c. bath. After renewal of the bath,
the addition of an identical amount of albumin induced no change
in the tone of the horns; however, upon the addition of barium
chloride a marked contraction resulted. None of the horns from the
negative cats showed any contraction except when barium chloride
was added to the bath.

Discussion

By correlation of the j^hysiological responses of the ])assively sensi-
tized cat to an intravenous injection of the homologous antigen, it is
possible to describe anaphylactic shock in the anesthetized animal.
Soon after the intravenous injection of the shock dose, the arterial
blood pressure abruptly falls, and at the same time the heart rate is
markedly reduced. The oncometric kidney volume is also greatly
diminished. The intestinal pressure definitely increases and at times
this is also true of the intracystic pressure. Marked specific contrac-
tions are exhibited by excised intestinal strips and uterine horns
when brought in contact with small amounts of the homologous pro-
tein. The coagulation time of the blood and the rectal temperature
are apparently unaffected. This would seem to indicate that a hep-
arin-like substance is not liberated during anaphylactic shock in
the cat. There are no demonstrable circulating antibodies for crys-
talline egg albumin ])resent. This fact would indicate that the anti-
bodies injected are either quickly fixed by the animal's tissues or
hastily excreted. Since anaphylactic shock could be demonstrated,
the former suggestion seems more fitting. There is 100 percent cor-

Kabler and Sherwood: Studies in Anaphylaxis 167

relation of the excised smooth muscle responses with those of the
arterial blood pressure, kidney volume, etc.

It has been i^ointcd out that all the rabbit antisera used for passive
sensitization in these studies were capable of passively sensitizing
guinea pigs to a subsequent injection of crystalline egg albumin, yet
only forty percent of tlie cats in this series were rendered hypersensi-
tive to the same antiu:t'ii.

The guinea pig has long been considered the animal "par excel-
lent" for demonstrating the anaphylactic phenomena. It has been
shown by Spain and Grove (8) , Sherwood and Stoland [9) , Tim and
Kuratchkin ilO) and others that the titer of the anaphylactic sensi-
tizer did not parallel the titer of the i)recipitating antibody in anti-
sera. Our results in jiassively sensitizing cats with rabbit antisera
bear out these findings. It is likewise demonstrated that the cat
does not yield to i)assive sensitization as readily as does the guinea
pig. Sherwood and Stoland (10) have shown that some dogs are not
passively sensitized, while others are by injections of the same anti-
serum. This recalls to mind the additional factor of individual
variation.

Although the physiological responses of the cat to histamine and
peptone injections resemble those of anaphylactic shock in the cat,
there are sufficient differences to render it questionable that either
histamine or j^eptone is solely responsible for the anaphylactic phe-
nomenon. One of us, Sherwood ill), has recently summarized and
discussed the evidence for and against histamine as the essential
factor in anaphylactic shock.

Conclusions

From the results of the foregoing jjliysiological studies the follow-
ing conclusions were drawn:

1. That in the light of our results, it ai)i)ears that the so-called
anaphylactic i)henomena in cats as reported by earlier investi-
gators were exaggerated "Brodie Reactions."

2. That active sensitization does not occur in cats, or is so rare
that it was not observed in the series of this study.

3. That the refractory character of cats is due to an inability to
generate sufficient antibodies, since only slight antibody for-
mation was demonstrated following the injection of vastly dif-
ferent antigens such as sheep cells, ciystalline egg albumin and
suspensions of bacteria.

4. That cats can be passively sensitized by injecting intraperi-
toneally adequate amounts of high-titered rabbit antisera.

168 The University Science Bulletin

5. That the anaphylactic responses of such passively sensitized
cats are specific, and that the cats show complete desensitiza-
tion upon reinjection of the specific antigen.

6. That the Dale reaction can be used as a criterion of feline sen-
sitization.

7. That the symptoms of experimental anaphylaxis in the anes-
thetized cat are:

i. Profound drop in blood pressure.

ii. Marked decrease in heart rate.

iii. Decided decrease in kidney volume.

iv. Increase of intestinal pressure in the majority of ani-
mals.

V. Specific contractions of excised smooth muscle such as
intestinal strips and uterine horns.

vi. Desensitization upon reinject'on.

BIBLIOGRAPHY

1. Brodie. J. of Physiol. 1900, 26. 48.

2. Manwahino, W. H. Zoit. fiir Imimmil., 1910, 8. 1.

3. ScHULTz, W. H. J. of Pharm. and E.xper. Theiap., 1911, 3, 299.

4. Edmunds, C. W. Zeit. fiir Immunit.. 1914, 22, 181.

5. Drinker, C. K., and Bronfenbrenner, J. J. of Imnuuiol., 1924, 9, 395.

6. Kahler, P. Univ. of Kansas Science Bulletin, 1938, 25.

7. Hopkins, F. G., and Penkus, S. X. J. of Physiol.. 1898, 23, 130.

8. Sp.ain. W. C, and Grove, E. F. J. of Imniiinol.. 1925, 10, 433.

9. Sherwood. N. P., and Stoland, O. O. J. of Immunol., 1930, 20, 101.

10. Tim, C. E., and Kiratchkin, J. J. Pioc. Sov. Exp. Biol, and Med., 1932,
29, 1151.

11. Sherwood, X. P. Imnuniology, C. J. Mosby Co., St. Louis, 1935, 485-490.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV I June 1, 19.3S [No. 6

A New Fish, Listracanthus eliasi, from the Pennsyl-
vanian of Nodaway County, Missouri

CLAUDE W. HIBBARD,

Muse'im of \'erteliiate Paleontoloax'. Univer.'iity of Kansas, LawifUce

Abstr.\ct: Listracfuithus eliasi Hibbaid n. sp., ba.sed on complete spine and
two associated spines, smallest known species of the genus Listracanthus.

AMONG some ichthyodoriilites in the Museum of Vertebrate
Paleontology, collected by Prof. M. K. Elias of the Kansas
Geological Survey, are three spines representing an undcscribed
form of ListracanfJtus.

Listracanthus eliasi n. sp.

(P!. XIX)

Types: Holotyiie No. 884F, University of Kansas Museum of
Vertebrate Paleontology. Comj^lete spine with truncated base.
Paratypes, K.U.AI.V.P. 885F-6F; dorsal part of spines lacking base.

Horizon and Type Locality: Taken from a concretion a few feet
above Elmo coal, in the Ccdarvale shale, Wabaunsee group, Virgil
series of the ^Middle Pennsylvanian of Mid-Continent. Taken by
M. K. Elias, 1934, at Carpenter's coal mine, 2 miles west and 3
miles soutli of Burlington Junction, Nodaway county, IVIissouri.

Diagnosis: The sjune is delicate, flattened and \'ery thin. The
sides contain from nine to ten longitudinal carinae at the base of
the spine. C)nly five longitudinal carinae continue throughout the
length of the spine, these being the five nearest the convex edge.
The outermost carinae on the concave side of the spine at its base
are lost as the spine gradually tapers off from the base to its apex.
The concave side of the sjnne is set with slender, sharply pointed,
closely packed teeth which are directed upward. The teeth are

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170 The University Science Bulletin

missing from the convex side with the exception of the extreme tip
of the spine where a few scattered teeth may occur. The spine
narrows abruptly above the truncated base on the concave side,
giving a gentle arched appearance. The holotype is 15 mm. in
length; width of truncated base, 3 mm.; greatest width of spine
•above base, 2 mm.; the teeth first appear on the concave side of
the spine 4 mm. from the base, on the convex side, 13 mm. Nine
longitudinal carinae are present with only five extending to the tip
of the spine. The paratypes agree with the holotype except that the
bases of the spines are missing. No. 886F is 15 mm. long, liaving ten
longitudinal carinae, only five continuing to tip of the sjiine. No.
885F with base missing is 15.5 mm. in leng-th and ])ossessing ten
longitudinal carinae.

Listracanthus eliasi may be distinguished from the other forms
by its small size and the fact that the teeth are nearly missing on
the convex surface and confined only to the tip of the spine. Though
Listracanthus has been found m many localities in Europe it is
one of the rarer forms found in the Pennsylvanian of North America,
being known only from the coal measure of Ohio and Illinois; also
from a single locality in Andrew county, Missouri, in the Cherokee
shale just overlying the Springfield coal.

Hibbard: a New Fish

171

PLATE XIX

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXV I June 1, 1938 [No. 7

Distribution of the Genus Reithrodo7itomys in Kansas

CLAUDE W. HIBBARD,
Museum of Vertebrate Paleontology, University of Kansas, Lawrence

Abstract: The occurrence of Reilhrodontoynys albescens albescens (new
state record); R. a. griseus; B. megalotis clychei and R. julvescens aiirandus
in Kansas with notes on distribution and habits.

npHE purpose of this paper is to give the distribution of the har-
J- vest mice of Kansas as represented by the specimens in the
Kansas University Museum of Mammals. In no way does it present
a clear picture of the distribution within the state due to the many
areas not yet studied. Field notes are included wliich may be of
value to others collecting in the field.

In the past six years of collecting in Kansas the following obser-
vations have been made on the habitat of Peromyscus and Eeithro-
dontomys. The genus Peromyscus occurs abundantly through the
state in all counties. In all of the counties carefully worked both
the Maniculatus and Leucopus groups have been found. Though
both occur in the same region, they inhabit entirely different ecologi-
cal stations. Peromyscus leucopus is confined chiefly to the more
hinnid areas of a given region; that is, along streams and lowlands.
Peromyscus manicidatus is found in the drier parts of a region of
upland meadows, fields and pastures. There is a slight overlapping
of range in the eastern one third of Kansas, that area east of the
Flint Hills known as the Osage Plains. After passing west of the
Flint Hills it is found that Peromyscus leucopus never encroaches
upon the areas inhabited by P. mainculatus, but is restricted entirely
to the most humid stations, especially along the more permanent
streams. It is in this region that we find the Maniculatus group to
some extent encroaching upon tlie stations favored by leucopus. A

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174 The University Science Bulletin

slight increase in elevation of a few feet giving a strictly grassland
area is found to be a perfect barrier to leucopus upon entering the
range of maniculatus. It is probable that the few maniculatus taken
along the streams with leucopns in the extreme western part of the
state come to these stations for water.

The Albescens group, trapi^ed in Greenwood, Douglas, Pottawa-
tomie, Barber and Hamilton counties, has been taken only in the
areas inhabited by Pei-omyscus maniculatus. The specimens of
Reithrodontom.ys m. dychei taken in Douglas county have been
found only to inhabit the humid stations occupied by Pcromyscus
leucopus noveboracensis. The nineteen specimens of R. m. dychei
taken in Rawlins county along Beaver creek in the summer of 1936
were found associated with Peromyscus leucopus aiidulus. Though
only a very few specimens of P. m. nebrascensis were taken in the
timber along the streams in association wdth P. I. aridulus and R.
tn. dychei, the latter was never taken away from the stream in the
surrounding uplands inhabited by P. m. nebi^ascensis. Insufficient
field work has been done in Kansas to state definitely that the ^4/-
bescens group inhabited only the stations occupied by Peromyscus
manicidatus, and that the Megalotis group will inhabit only the
stations occupied by Peromyscus leucopus. Though all evidence,
however, seems to point to this condition and it should be expected,
especially in the regions where both species of Reithrodontomys
occur.

Reithrodontomys albescens albescens Gary

1903. Rcithrtidohtomya albtscens Cary, Pioc. Biol. Soc. Washington, Vol. 16, p. 53.

Type Locality. Eighteen miles northwest of Kennedy, Gherry
county, Nebraska.

Range in Kansas. Hamilton, Kearny and Alorton counties.

Characters. More pallid with a greater yellowish wash than
griseus.

Measurements. In millimeters, of two adult males from two
miles east of Coolidge, Kan., Hamilton coimty; length, 120-121; tail,
45-54; hindfoot, 15-15; ear, 12-12; greatest length of skull, 19.3-
broken; zygomatic breadth, 10.1-10.3, respectively.

Remarks. Heretofore the range of Reithrodontomys a. albescens
Cary has been confined to the sand-hill region of Nebraska and
western Soutli Dakota; west to Loveland, Golo.

Nine s]:)ecjnens from Hamilton and Morton counties were found
to represent an undetermined race in Kansas. The specimens were
sent to A. H. Howell for identification and are referred to R. a.

Hibbard: Reithrodontomys in Kansas 175

albescens. It is not known under what conditions the specimens
were taken in ]Morton and Kearny counties, but the specimens
from Hamilton county were taken about two miles east of Coolidge,
Kan., on the north side of the Arkansas river. The harvest mice
were found on the secondary flood plain which supjjorts what is
called "Alkali or Salt Grass," in that area. The sand hills are south
of the river two or three miles. The specimens were trapped by
using rolled oats for bait. They were found under a different en-
vironment than those from Nebraska, South Dakota and Colorado,
and were taken in association with Peromyscus maniculatus. There
is a male specimen in the collection taken eight miles east of Keys,
Cimarron county, Oklahoma, July 8, 1932, by Eleanor Henderson
referable to this form.

The specimens are lighter than typical E. a. albescens and will
probably be found to represent a new race when sufficient material
is at hand; since they do not compare with R. montanus.

Reithrodontomys albescens griseus (Bailey)

190.5. Reithrodontomys griseus Bailey. North .\iiiciic;ui Fauna. No. 2,5, p. 106.

1914. Reitlirodontomys albescens yriseus Howell, Xoith .American Fauna, No. .30. p. 2.S.

Type Locality. San Antonio, Bexar county, Texas.

Range in Kansas. See map.

Characters. Similar to albescens, but darker; total length less
than 130 mm. and tail less than 60 mm.

Measurements. Average and extreme measurements in milli-
meters of 9 adult females from Kansas: length, 121 (108-126);
tail, 53 (50-55); hind foot, 15.6 (15-161; ear, 12 (12-12); greatest
length of skull, 20.1 (19. 1-21.41; zygomatic width, 10.6 (9.9-11.1).

Measurements of 10 adult males from Kansas: length, 118.5
(108-122) ; tail, 54 (44-561 ; hind foot, 16 (14-181 ; ear, 12 (11-13) ;
greatest length of skull, 20.4 ( 1!).1-21.7) ; zygomatic width, 10.6
(10-11).

Remarks. Reithrodontomys albescens griseus occupies the largest
range in Kansas of any of the harvest mice. It has not been found
abundant in any area, though it can be considered common. In the
C(!llccfon there are tliirty-seven specimens. Those that have been
trapped were taken by using rolled oats for Ijait. The large number
of Peromyscus maniculatus occurring in the same area probably
accounts for the fact that so few are taken by sets. In no I'egion
have we collected out the more abundant forms giving the less
abundant forms a chance to visit the sets. It is of interest that a
number of specimens have been taken at night while collecting

176 The University Science Bulletin

with a lantern in grassy areas. A niunber of specimens have been
taken along high, stony ridges in pastures while turning stones for
reptiles and amphibians.

Two immature specimens 83 mm. and 84 mm., respectively, were
taken July 10, 1911, in Clark county; a female was taken in Doni-
phan county, November 26, 1924, that contained 5 embiyos. During
April, 1932, a female (No. 8660 K. U. M. M.) was taken in Douglas
county while collecting at night and placed in a cage in the labora-
tory. She became very tame and allowed herself to be handled.
A nest was built out of cut bluegrass in which 5 young were born,
April 25. Four of the young lived in the laboratory until they were
made into study skins.

Reithrodontomys megalotis dijchci (Allen)

189.5. Reithrndoiitomyx dychei Allen. Bull. American Miis. Nat. Hist., \o\. 7, p. 120.

189.T. Rcithrodoutornyti dychei twbrasceiisis Allen. Bull. Anier. Mus. Nat. Hist., Vol. 7,
p. 122.

1914. ReithrodotitomyK mefialotis dychei Howell, North Anier. Fainia. No. 36, p. 30.

Type Locality. Lawrence, Douglas county, Kansas.

Range in Kansas. See map.

Characters. Distinguished from Albescens group l)y larger size,
more intense ochraceous coloration, lack of dark dorsal area, an oval-
shaped foramen magnum which is more rounded in Albescens group.
Length 130 mm. to 150 mm.; tail, 62 to 76 mm.

Measurements. Average and extreme measurements in millimeters
of 9 adult females from Kansas: length, 139 (130-149) ; tail, 66 (62-
711; hind foot, 17 (16-18); ear, 12.75 (12-13); greatest length of
skull, 20.8 (20-21.8) ; zygomatic width, 10.8 (10.3-12). Average of
6 adult males from Kansas: length, 140 (132-145) ; tail, 67 (63-76) ;
hind foot. 17 (16.5-17.5) ; ear, 13 (13-14) ; greatest length of skull,
21.3 (20.5-22.5) ; zygomatic width, 11.1 (10.5-12).

Reithrodontomys megalotis dychei, so far as known, is confined to
the northern one half of the state. It is found along the streams and
lowland fields. In Douglas county, where a more intensive study of
the mammals has been carried on than in any other county of the
state, fifty-three specimens have been taken, while only twenty-one
specimens of R. a. griseus were taken over the same i^eriod. The
ratio of Peromyscus leucopus to that of P. maniculatus taken in
Douglas county is considerably greater, 4:1 ratio. The specimens
of Reithrodontomys m. dychei taken in Rawlins county, in the eco-
logical stations found inlialuted by Peromyscus I. ardilus. live at a
considerably higher and drier ahitude than can be found on the
extreme uplands of Douglas county, though the stations inhabited

Hibbard: Reithrodontomys in Kansas 177

in Rawlins county are the most humid in that county. So far in
Douglas county only a slight overla])]nng has been found in the
range of the two species of harvest mice, though no greater than
that of the species of Peroniyscus.

A female, No. 3806, was taken March 28, 1920, in Douglas county,
which contains 4 embryos; female No. 6157, taken Ai)ril 26, 1928, in
Douglas county, contained 5 embryos 2 mm. in length. Two im-
mature specimens were taken July 28, 1896, in Leavenworth county,
in a timothy meadow, which measured respectively 82.5 mm. and
86.2 mm.

Reithrodontomys fulvescens auranthis (Allen)

ISflf). Reithrodoritoinyti ntexicatius aurUHtiuf: Allen, Bull. AiutM'. Muk. Nat. Hist., Vol.
7. p. 137.

1899. Reithrodontomys chrysotis Elliot, Field Colunili. .Mu.^^. P\il>l. 87, Zool. Ser., Vol.
1. p. 281.

1914. Reitlirodontomys fulvescevs auraiitiiis Howell. Noith .\iiier. Fauna, No. 36, p. 48.

Type Locality. Lafayette parish, Louisiana.

Range in Kansas. Confined to the Austrorii)arian zone of Kansas
known as the Cherokee lowland (see map).

Remarks. Known only from 2 specimens collected in Cherokee
county, respectively December 1913, and December 1915, by Vic
Housholder.

Specimen No. 4478, University of Kansas Collection, reported by
J. D. Black, Journal of Mammalogy, Vol. 16, No. 3, pp. 231-232, as
Reithrodontomys f. avrantius from Anderson county, Kansas, is the
skin of R. a. griseus. The specimen has been identified by Doctor
Kellogg of the U. S. National Museum. The measurements are typi-
cally those of R. a. griseus. The upper part and sides of the speci-
men are more richly colored and the underpart more heavily washed
with yellow than in tyjiical specimens of R. a. griseus, but correspond
with the color phase taken in Harvey county.

12—4141

178 The University Science Bulletin

PLATE XX

Map of Kansas showing the distribution of the Harvest Mice.
□ Reithrodontomys albescens albescens Gary.
O Reithrodontomys albescens grisens (Bailey).
A Reithrodontomys megalotis dychei (Allen).
V Reithrodontomys fidvescens auranlius (Allen).

Hibbard: Reithrodoxtomys in Kansas

179

PLATE XX

THE UNIVEESITY OP KANSAS

SCIENCE BULLETIN

Vol. XXV 1 June 1, 1938 [No. 8

A Lower Jaw of Martinogale alveodens Hall

By DAM I) HOSBROOK Dl'NKLE,
Biological Lalioratories, Harvard University

IN THE spring of 1935 Dr. Edward H. Taylor and the author,
on a field trip to western Kansas, were fortunate in securing the
riglit mandible of the little-known miistelid, Martinogale alveodens
Hall. The specimen was collected in the Lower Pliocene beds
of the SW14 sec. 26, T. 10, R. 38 W., Sherman county, Kansas. It
is sufficiently complete to warrant this further description.

I am grateful to Professors E. H. Taylor and H. H. Lane for
their help in the ]ireparation of this report; to Mr. C. D. Bunker and
Dr. Glover Allen for the use of recent material for comjiarison; and
to Mr. L. L Price for retouching the photographs.

This specimen (Kansas Univ. Mus. Vert. Pal., No. 3833) con-
sists of a complete right mandible with C, Pm^, and M^. The re-
mainder of the teetli are represented only by their alveoli. It is
undoubtedly the jaw of a young adult, as the teeth are practically
unworn.

The mandible of Martinogale alveodens is about two thirds as
long as that of Spilogale interrupta and is quite similar in general
contours. The inferior margin does not exhibit the marked antero-
posterior convexity of profile seen in the mandible of Mustela
cicognanii cicognanii and the jaw is less heavily constructed. A dis-
tinct chin is develoi)ed anteriorly, a feature faintly discernible in
Spilogale but lacking in Mustela. The symphysis, in contrast to its
inferior posterior elongation in living weasels, is restricted ]:)0.steriorly
and is no broader inferiorly than the maximum depth. The three
mental foramina present are situated laterally below the canine,
below Pm., and below Pm^, respectively. The number and j^osition
of the mental foramina are to be considered as variable characters
as only two were i)resent in the type specimen; in all specimens of

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182 The University Science Bulletin

Spilogale examined the number varies from 2 to 4, and from riglit
side to left side in widely divergent positions. The coronoid process
shows a development proportionately similar to that of recent
skunks, except that in side view the posterior face has a slightly
more concave profile. The articular process is relatively more
heavily developed. The masseteric fossa is less strongly pronounced
and the angle of the jaw is more prominent than in Spilogale.
Mustcla differs from Martinogale in the above respects, mainly in
the convex profile of the posterior face of the coronoid process in
side view, and in the greatly receding angle of the jaw.

The dental formula in the mandible is I^ ^ 3. Cj, Pnu ^ ^, and M^^„.

The incisors, judging from the alveoli, are crowded together, so that
they lie in a jilane nearly parallel to that of the symphysis instead of
transversely. The relationships of the incisors are the same as those
found in Spilogale interrutpa and Mustela cicognanii cicognanii.
The only difference apparent is the greater degree of mesial crowd-
ing and consequent greater reduction of I^ found in Martinogale.
The alveolus of L, is large and is located between the symphysis and
canine. Directlv beneath is the much reduced alveolus of L. The
alveolus of I3 lies almost directly below but a little lateral to tliat
of I.J and closely approximates it in size.

The canine is proportionately similar in length and contours to
that of Spilogale interrupta. Both differ from Mustela cicognanii
cicognanii in sharper upward curvature and in a decidedly more
prominent cingulum on the posterior and medial surfaces. The
alveoli of Pnio indicate that it was a small double-rooted tooth
crowded forward against the base of the canine so that it sat trans-
versely in the jaw. The alveoli of Pm^ are somewhat larger than
those of Pnio and lie laterally on the superior surface of the mandi-
ble, indicating that the tooth had its anterior end crowded outward,
forming nearly a right angle with Pm^. Pm^ is large with no acces-
sory cusps, but a well-developed heel and a smaller cusp anteriorly
on the lingual side. The main cusp of Pm^ is produced upward and
backward and the tip is higher than the protoconid of ^NIj. A cin-
gulum extends entirely around tlie tooth, and is more pronounced
medially. Three sharp crests, anterior, medial, and posterior, ex-
tend from the tip of the crown to the cingulum.

The lengtli of ]\I, is equal to its distance from the canine. The
lirotoconid is higher tlian the paraconid. The metaconid, located
directly medial to the protoconid, is but slightly shorter than the
paraconid and hidden by it in side view. The lateral crest of the

Dunkle: Martinogale Alveodens 183

lieavily basined talonid is carried medially behind the protoconid
and is separated from the posterior crest of the latter by a distinct
notch. The sharp and distinct medial crest of the talonid is not as
high nor as heavily developed as the lateral crest. It is continuous
with the posterior crest of the metaconid. An indistinct hypoconid
is apparent on the lateral crest. A weakly developed cingulum may
be traced around the entire tooth. The alveoli of Mo show that this
tooth was less reduced than in the living forms of the subgenus
Mustcla, and was double rooted.

Comparison with the type material shows the specimens to be
similar in most respects. The chief differences are listed below.

(1) On Mj there is a distinct notch between the lateral crest of
the talonid and the posterior crest of the protoconid, whereas the
corresponding crest of the talonid in the type was described as being
continuous with the posterior crest of the protoconid.

(2) The alveoli of Pm. indicate a smaller double-rooted tooth.
The single alveolus of Pm^ in the type is larger than the alveoli of
Pm.,.

(3) The indices of Pm^ and M^ (width divided by length) are
0.64 and 0.47, respectively, as compared with 0.50 and 0.38 of the
corresponding teeth of the type and thus indicate proportionately
shorter and broader teeth.

These differences may be explained on the basis of sexual di-
morphism in view of the great number of striking similarities be-
tween the two specimens and the improbability that two closely
related species would be found in a deposit of such limited local ex-
tent. This is especially probable since sexual differences in size and
to lesser degree in structure are markedly evident in the living rep-
resentatives of this group.

Summarizing, Martinogale differs from Mustela not only in the
general contours of the mandible: the distinct chin, short deep
symphysis, the concave posterior profile of the coronoid process, and
the prominent angle of the jaw; but also in dental characters: the
crowded Pm., in process of reduction, the extremely strong meta-
conid, the well-developed lateral crest of the heavily basined talonid
which exhibits no tendency toward a developing trenchant condition,
and in the less reduced and still double-rotted M.. It is of jiar-
ticular interest to note that all these differences from Mustela de-
note points of similarity between Martinogale and Spilogale, as com-
parisons throughout the above descriptions show. On the basis of
these osteological and dental characteis this Pliocene species should

184 The University Science Bulletin

be classed with Spilogalc until the discovery of the skull shall indi-
cate its true phylogenetic position.

Measurements of Specimens of Martinogale alveodens Hall in the
University of Kansas Museum of Vertebrate Paleontology

Xn. 3473 No. 3833

(Type)

MM. MM

Total length of jaw 21.9

Length of tooth row from posterior base of C

to posterior base of Mi 9.9

Length Pm3 2.5 2.0*

Breadth Pms 1.0 1.3*

Length Pn^ 3.0 2.8

Breadth Pm4 1.5 1.8

Length Pm2 to and including Pni4 5.0*

Length Mi 5.8 5.3

Transver.se breadth at metaconid 2.1 2.5

Length of talonid 2.5 2.1

Breadth of talonid 2.0 2.1

Depth of mandible at anterior base of Mi 3.3 3.4

Breadth of mandible below trigonid of Ml 2.2 2.2

* Measurement of Alveoli.

REFERENCES

Elw.s. Maxim K. 1931. The Geology of Wallace Countv, Kansas. Bull.
State Geol. Surv. Kansast. Xo. 18, pp. 1-254, Pis. 1-42.

H-ALL, E. R.^YMOND. 1930. Three New Genera of Mustelidae from the Later
Tertiary of North America. J. Mamm., Vol. 11, No. 2, pp. 146-155, Pis. 7-8.

Dunkle: Martinociale Alveodens

185

m^

•^yf

/.

EXPLANATION OF PLATE XXI

Figs. 1 to 3. Right mandibular ramus and lower teeth of Mdiiiiiof/ale alveo-
dens Hall (1930), No. 3833, Univ. Kan. Mus. Vert. Paleontology; from sand
deposit near the top of the Lower Pliocene, SW V^, sec. 26, T. 10, R. 38 W.,
Sherman county, Kan.sas.

1. Crown view, X 4.

2. Medial view. X 4.

3. Lateral y'\e\\, X 4.

;

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1,1988 [No. 9

Weights and Linear Dimensions of the Skull and of Some
of the Long Bones of the Mourning Dove {Zenaidura
macroura carolinensis)

HOMER B. LATIMER and C. WILLET ASLING,
Department of Anatomy, University of Kansas

A CAREFUL search of the literature shows that the number of
statistical studies on bird skeletons is just about as limited as
it is for mammalian skeletons. Schneider and Dunn ('24) studied
the length and breadth of the skull and the lengths of four long
bones in a series of 350 adult male and 46 adult female chickens.
The length and width of the head and the weights and lengths of
the six long bones of the turkey hen have been reported (Latimer
and Rosenbaum, '26). Lerner ('37) has reviewed the literature on
the growth of the chicken skeleton, but for the adult forms of other
birds there is very little except for a few measurements in the litera-
ture on taxonomy. This series of dove skeletons is not as large as
could be desired, but it is hoped that these data will be of some
value in filling a gap in our knowledge of avian osteology.

Following the plan of the two earlier papers on the mammal
skeletons, this study will give first the average weights and dimen-
sions of the skull and of six of the long bones, the coefficients of
variability and the sex differences in the various measurements.
Next the correlations between the skull weight and the weights of
the long bones and the correlations between three dimensions of
the skull with the other skull dimensions and the lengths of the long
bones will be presented. Very frequently linear dimensions of the
skull are given in taxonomic papers and this study of the relative
lengths should show how valid the use of these measurements is in
giving an idea of tlie size of the other parts of the skeleton. The
last topic will be tlie question of symmetry of the paired bones.

Material and Methods

These skeletons are a part of the University Museum collections
and they were all prepared in the same excellent manner. The

(187)

188 The University Science Bulletin

inothocl is the same as tliat used in tlic jireparation of the earlier
reports on the other two series of skeletons. A very few of the parts
were injured so that weights or measurements could not be made,
l)ut as many measurements as possible were made on each skeleton.
One of tlie skeletons was evidently that of a young bird with in-
complete ossification, and tliis entire skeleton was not used in this
study. There were 26 male and 14 female skeletons used. The
small numbers of each sex, especially the females, will not warrant
final conclusions as to sex differences.

The weights were made first, or before the bones were handled
much. The bones were not oven dried, but the bones of an entire
skeleton were kept together in a small pasteboard box and the boxes
were stored in a dry room for some time before weighing. Each of
the paired bones was weighed and measured separately and the
weights an<l lengths in all but table four are the sum of the weights
or the average length of the two bones. In making the weights as
well as in measuring the lengths, the first one of a pair of bones
picked up was weighed or measured, so that there might not be
constant sequence and possible personal eciuation in these measure-
ments. All weights were made on a chemical balance sensitive to
\/jo milligrauL The bones were not weighed in closed containers
and the weights were recorded to the nearest milligram.

A vernier caliper reading to \/n, millimeter was used in making all
of the linear measurements, except the total length and wingspread.
These two measurements were recorded on the data card with each
skeleton and were made at the time the skeleton was prepared by the
museum workers. The wingspread was given for but ten skeletons,
equally divided as to sex. The methods of making ten of the
skull measurements are indicated in figure 1. The skull height is
the maximum height with the fixed ami of the caliper held parallel
to the base of the skull. The length of the mandible is the maximum
length from the symphysis to the posterior end of the left side. The
lengths of the long bones are all the maximum length with the long
axis of the bone held parallel to the back of the caliper. The sternal
length is the maximum lengtli in a line connecting the anterior end
of the keel and the posterior end held parallel to the back of the
caliper. The sternal height is the maximum depth of the keel meas-
ured from the inside of the sternum just posterior to the anterior end
and with the anterior border of the keel held parallel to the back of
the caliper.

The formulae used in computing the values given in the tables are
those given by Dunn ('29). All of the computations were carried to

Latimer and Asling: The Mourning Dove

189

more decimal ])laces than given in the tables and all computations
were carefully checked. The junior author made all of the linear
measurements and the weighing and the preparation of the paper
were done by the senior author.

We wish to express our gratitude to Mr. C. D. Bunker for per-
mitting us to study these beautifully prepared skeletons and to Miss
Edna Mae McConnell for her careful work in aiding with the sta-
tistical work.

Measurements, Variation and Sex Differences

The average ponderal and linear measurements and their probable
errors are given in the second column of table 1. The last column
gives the significant ratios or the difference between the measure-
ments of male and female divided by the probable error of the differ-
ence. The least variable of the coefficients of variation of the

Table 1

Average measurements, coefficients of \uriation and significant ratios

Skull and mandible

Humervus

Radius

Ulna

Femur

Tibiofibula

Tarsometatarsus

Skull length

Basion-oci-iput

Basion-spine sphenoid . . .
Spine sphenoid-tip beak.

Occiput-hinge

Hinge-tip beak

Mandible

Skull height

Bitemporal width

Interorbital width

Foramen magnum lensth
Foramen magnum width

Sternal length

Sternal height

Interacetabular width . . .

Humerus

Radius

Ulna

Femur

Tibiofibula

Tarsometatarsus

Total length

Wingspread*

Average

Coe

IHcient

Difference

weight in
grams.

of
lat

var

on.

P. E. diff.

0.345

± 0.0035

9.42

±

0.73

2.10

0 . 527

=t 0.0033

5.91

±

0.45

3.73

0.165

=t 0.0018

9.96

=t

0.76

1.70

0.399

± 0.0054

12.29

±

0.97

0.80

0.240

± 0,0045

17.38

±

1.37

3.48

0.323

± 0.0037

10.50

±

0.82

0.84

0.138

± 0.0015

10.24

±

0.79

4.71

Average length

in millimeters.

41 .64

± 0,191

4.29

zt

0.32

1.22

7.06

± 0.058

7.57

±

0.58

0.15

8.13

± 0.038

4.28

±

0.33

4.33

27.14

± 0,108

3.69

±

0.28

0.61

26.13

± 0,087

3.12

zfc

0.24

3.10

17.67

=t 0.100

5.32

±

0.40

1.11

29.74

=t 0.129

3.92

±

0.31

0.89

15.80

± 0.063

3.63

=t

0.28

1.00

15.95

± 0.039

2.30

:±z

0.17

3.02

9.02

=t 0.093

9.69

zt

0.74

1.13

3.53

=t 0.031

7.80

±

0.62

1.61

4.36

± 0.024

5.19

±

0.39

0.09

52.52

=t 0.213

3.66

±

0.29

5.34

20.42

± 0.092

4.19

±

0.32

4.07

18.73

± 0.103

5.07

zt

0..39

0.52

31.98

=fc 0.079

2.30

±

0.17

3.24

33.40

=t= 0.109

3.06

±

0.23

2.88

36.87

± 0.102

2.. 52

zfc

0.19

2.81

28.35

± 0.083

2.70

=fc:

0.21

3.97

38.01

± 0.111

2.68

±

0.21

3.68

20.91

± 0.075

3.34

±

0.26

4.56

280.73

± 2.091

6.71

=iz

0.53

2.99

148.45

± 2.17

6.84

zfc

1.04

* Ten cases.

190

The University Science Bulletin

weights is the humerus, with the skull weight second. The weight of
the femur is the most variable, or 2.94 times as variable as the
weight of the humerus. The linear dimensions are less variable, as
is to be expected, and the humerus and the bitemporal width of the
skull have the lowest coefficients of variation. The three bones of
the hind limb are 36 percent more variable in weight and 11 percent
more variable in length than the three bones of the fore limb. The
interacetabular diameter and the two dimensions of the sternum are
more variable than the lengths of the six long bones and the total
length and the wingspread are still more variable. The average of
the coefficients of variation of the six long bones is 2.77 percent and
the similar average for the twelve dimensions of the skull is 5.07
percent.

The bitemporal width or skull width is the most constant of all
of the skull dimensions, and it happens to be the same as the lowest
coefficient of variation of the lengths of the long bones, or 2.30 jier-
cent. The interorbital diameter is the most variable of all of the
skull dimensions, thus these two transverse measurements form the
two extremes of all of the skull dimensions. The interorbital di-
ameter is limited by a thin edge of bone on each side as shown in
figure 1. The central part of the bone provides adequate strength
and so the edges may vary considerably without any significant re-

T.\BLE 2

Average measurements and coefficients of variation for males and females

Males.

Fema

les.

Average weight
in grams.

Coefficient
of variation.

Average weight
in grams.

Coefficient
of variation.

Humerus

0.535 ± 0.0038
0.227 ± 0.0046
0.143 ± 0.0019

5.41

14.71

9.70

± 0.51
± 1.46
±0.95

0.511 ± 0.0051
0.262 ± 0.0089
0.130 ± 0.0021

5.32

18.25

8.60

±

0.71

Femur

2 49

1 .15

Average length
millimeters.

Average length
millimeters.

Humerus .

32.19 ± 0.07
28.60 ± 0.08
38.35 ± 0.10
21.19 ± 0.06

8.27 ± 0.03
26.32 ± 0.11

16.05 ± 0.04

.53.22 ± 0.27
20.65 ± 0.12

1.62
1.93
1.93
2.12

2.88
3.07

1.77

3 . 55
4.24

± 0.15
± 0.19
± 0.19
± 0.21

± 0.28
± 0.29

± 0.17

± 0.35
± 0.40

31.59 ± 0.17
27.90 ± 0.16
37.43 ± 0.23
20.43 ± 0.16

7.91 ± 0.08
25.77 ± 0.14

15.77 ±0.08

51.24 ± 0.26
19.96 ± 0.12

2.94
3.04
3.26
4.07

5.14
2.86

2.81

2.60
3.11

±
±

zfc

±
±

0.39

Femur

0.40

Tihiofibula

0 44

Tarsometatarsus

0,54

Basion-spine sphenoid ....
Occiput-hinge

Bitemporal width

Sternal length

Sternal height

0.68
0.38

0.37

0.36
0.41

Latimer and Asling: The Mourning Dove 191

suit. On the other hand, the bitemporal diameter marks the maxi-
mum width of the brain case, and its width is much more significant.

In these skulls the bitemporal or skull width is the most constant
of the twelve dimensions of the skull. The skull height is next and
the skull length the most variable of these three. As shown in figure
1 (S. L.) the skull length includes the length of the brain case and
the length of the beak. The dimension which more nearly gives the
length of the brain case and is also comparable to the measurement
of Schneider and Dunn ('2-1) in the chicken skull is the occiput-hinge
dimension (0. H. in fig. 1) which has a coefficient of variation of
3.12 percent, or a lower coefficient than the skull height, but still
greater than the transverse or bitemporal diameter. Thus the greater
variability of the total skull length seems to be in the beak. The
dove skull with a more constant width, or bitemporal diameter, re-
sembles the skulls of the turkey (Latimer and Rosenbaum, '26),
chicken (Schneider and Dunn, '24) , muskrat (Latimer and Riley,
'34) and the mouse. Mm musculus (Green, '32). In the following
fonns the skull length has been found to be more constant: the
skunk (Latimer. '37), the mouse, Miis bactrianus (Green, '32) and
the human skull (Duckworth, '17; Orensteen, '15 and '20; Pearson
and Davin, '24, and others).

In general, the lengths of the six long bones are less variable than
the twelve linear dimensions of the skull. The skull width is less
varial)le than any of the other dimensions of the skull, with both the
skull height and length more variable and in this order. The aver-
age coefficient of variation for the twelve dimensions indicates that
the skull of the mourning dove is more variable than that of the
other bird skulls studied, and for the skunk (Latimer, '37), and yet,
it is less variable than for many mammal skulls.

The last column in table 1 gives the significant difference between
the weights and lengths in the male and female doves. This signifi-
cant ratio is the difference between the average measurements of
male and female divided by the probable error of this difference.
This shows that only three of the seven weights are significantly
heavier in the males, and nine of the twenty-two linear dimensions.
In eveiy case the weights and linear measurements were somewhat
greater in the males. No difference for the wingspread was de-
termined, for there were only five of each sex and it was felt that
this number was not large enough to warrant any conclusions. The
numbers of the specimens for these computations are not as large as
they should be, but the results would indicate that the mourning
do^'e does not show anv verv marked sex differences.

VJ2

The University Science Bulletin

T.ABLE 3

A. Correlations with skull weight. All weights in grams

Humerus

Radius

Ulna

Femur

Tibiofibula

Tarsometatarsus

+ 0.487
+ 0 . 544
+ 0.486
+ 0.322
+ 0.381
+ 0.619

0

082

0

076

0

085

0

098

0

095

0

068

B. Linear correlations. All lengths in millimeters

Skull length.

Skull height.

Bitemporal.

+ 0.155 ±0.104
-f 0.221 ± 0.101
+ 0.263 ± 0.102
+ 0.269 ± 0.100
+ 0.363 ± 0.095
+ 0.394 ± 0.091

— 0.431 ± 0.088
+ 0.185 ± 0.106
+ 0.368 ± 0.093
+ 0.325 ± 0.095
+ 0.369 ± 0.092
+ 0.481 ± 0.084

+ 0.157 ± 0.107
+ 0.1.52 ±0.104
+ 0.339 ± 0.094

— 0.126 ± 0.109

— 0.043 ± 0.106

+ 0.282 ±0.102
+ 0.323 ± 0.097
+ 0.104 ± 0.107

+ 0.381 ± 0.095
+ 0.6.59 ± 0.121

-h 0.296 ± 0.100
+ 0.317 ± 0.098
+ 0.230 ± 0.106
+ 0.308 ± 0.100
+ 0.310 ± 0.102
+ 0.376 ± 0.095

+ 0.021 ± 0.109
+ 0.342 ± 0.098
+ 0.183 ± 0.107
+ 0.345 ± 0.096
+ 0.073 ± 0.109
+ 0.275 ±0.105

+ 0 388 ± 0 091

Radius

Ulna

Femiir

-^ 0.546 ± 0.075
+ 0.657 ± 0.062
+ 0 492 ± 0 082

Tibiofibula

Tarsometatarsus

+ 0.576 ± 0.073
+ 0.477 ± 0.083

+ 0 085 ± 0 107

Basion-spine sphenoid

Spine sphenoid-tip beak

Occiput-hinge

Hinge-tip of beak

+ 0.4.50 ± 0.087
+ 0.770 ± 0.044
+ 0.529 ± 0.077
+ 0 086 ± 0 106

Mandible

+ 0 248 ± 0 104

Skull height

+ 0 304 ± 0 099

Bitemporal width

+ 0.304 ± 0.099
+ 0.393 ± 0.092

+ 0.274 ± 0.103
+ 0.008 ± 0.109

+ 0.316 ± 0.103
+ 0.171 ± 0.108
+ 0.272 ± 0.103

+ 0.318 ± 0.102
+ 0.463 ± 0.177

Interorbital width

Foramen magnum length

Foramen magnum width

Sternal length

Sternal height

Interacetabular width

+ 0.475 ± 0.083

— 0.088 ± 0.110
+ 0.232 ± 0.101

+ 0.365 ± 0.096
+ 0.435 ± 0.088
+ 0 545 ± 0 076

Total length

Wingspread

+ 0.447 ± 0.087
+ 0.168 ± 0.207

Table 2 gives the averages and the coefficients of variation for
males and females for those measurements in which there is in-
dicated a significant sex difference in table 1. In every case the
male measurements are greater than the female. In addition this
table shows that the females are more variable on the average. The
average coefficient of variation for all of these twelve measurements
is 4.41 percent for the males and 5.17 percent for the females or 17
percent more variable. The weights alone are 7.85 percent more
variable in the females and the linear dimensions are 28.79 percent
greater.

Correlations

In order to determine how well the weight and linear dimensions
of the skull represent the other measurements of the skeleton, the
skull weight was first correlated with the weights of the six long
bones and given in the first part of table 3. The correlations be-

Latimer and Asling: The Mourning Dove

193

tween the skull length and the other linear dimensions are given in
the first column of the second part of table 3. These were so low
that two other skull measurements, namely skull height and bitem-
])oral width or skull width, were correlated with the linear dimen-
sions and these correlations are given in the last two columns of the
second part of this table.

Table 4
Asymmetry in weight and length of the paired bones

The hea\ier or longer bone, or bones of equal siz;', are given in teuns of percentage of the
total number of each bone, which is given in the second column.

Weights.

Number

of

bones.

Right
greater.

Left
greater.

Same.

Humerus

Radius

Ulna

perniir

33
35
29
32
32
37

42.4
42.8
41.4
37.5
59.4
29.7

48.5
42.8
51.7
53.1
31.2
48.6

9.1

14.4

G.9

9.4

Tibiofibula

9.4

Xarsoniptatarsiis . .

21.7

Lengths.

Humerus

Radius

33
33
29
32
32
36

30.3
40.0
37.9
28.1
43.8
27.8

30.3
34.3

34.5
28.1
31.2
27.8

39.4
25.7

Ulna

27.6

43.8

Tibiofibula

25 . 0

Tarsometatarsus

44.4

The correlations of the weights of the six long bones with skull
weight give an average correlation of + 0.473. The similar average
correlation for the muskrat and the skunk both are above + 0.8.
We may conclude that the skull weight of the dove is not nearly as
good a criterion of the weights of the long bones as in the two species
of mammals. The average of the skull length correlated with all of
the linear dimensions is -f- 0.236, the similar correlation with the
skull height is + 0.266 and with the skull width, + 0.390. The
average correlations of the skull dimensions alone with the skull
length is + 0.161, with skull height, + 0.222 and with bitemporal
width, -|- 0.309. Thus it is evident that each of these dimensions of
the skull gives better average correlations with the lengths of the
six long bones and the other five dimensions than with the other
dimensions of the skull. The only one of these averages over -|- 0.5
is that with the skull width and lengths of the six long bones, or
+ 0.523. In the muskrat the average correlations of skull length
and the other dimensions of the skull and the lengths of the long
bones both averaged above + 0.7 and in the skunk the similar cor-
relations were slightly lower, but all far above the similar correla-

13—4141

194 The University Science Bulletin

tions in the mourning dove. Green ('33) finds some low and even
negative correlations with the skull length in the mouse. The wing-
spread is the only dimension which has a correlation above 0.5
percent when correlated with the skull length. Not one of the
twenty-one dimensions gives a correlation above 0.5 percent w^hen
they are correlated with skull height. When these dimensions are
correlated with bitemporal width there are six correlations above
0.5 percent. It has been shown above that the variability of the
dove, while it is greater on the average than that in some other
birds, is lower than that of both the skunk and the muskrat and
yet the correlations are much higher in both of these mammals.
This would lead to the conclusion that while there seems to be but
little variation in the same dimensions as measured in various doves,
the relative proportions of the dove are not as constant as in some
of the mammals.

In the skunk (Latimer, '37) the length of the foramen magnum
had a low correlation with skull length and so in this study both the
length and the width of the foramen magnum were measured, and as
shown in table 3 both of these dimensions when correlated with
either skull length, height, or width give very low correlations which
are not significant in any case. To see whether there was any uni-
formity in the tw^o diameters of the foramen magnum its length and
width were correlated with the resulting correlation of + 0.178 ±
0.107. This means that not only are the two dimensions of the fora-
men magnum not correlated with the three skull dimensions, but
that they bear little relationship to each other.

The length from the occiput to the anterior border of the foramen
magnum or basion-occiput (fig. 1) shows low correlations with all
three of the skull dimensions and so it was correlated with the
measurement from the basion to the tip of the spine of the sphenoid.
This should give some idea of the variability of the occipital con-
dyle with reference to the two ends of the basilar surface of the
skull. The correlation between the basion-occiput and the basion
to the spine of the sphenoid (fig. 1) was — 0.345 ±: 0.096, or a non-
significant negative correlation.

From the above correlations ^ve would judge that neither the
weight of the mourning dove skull, nor the skull length, skull
height, or the skull width are good criteria of the w^eights or lengths
of the long bones or of the other dimensions of the skull. Of the
three dimensions, the skull width or the bitemporal diameter is the
best criterion of the other linear measurements, and it is on the

Latimer and Asling: The Mourning Dove 195

average better correlated with the lengths of the long bones than
with the other skull dimensions.

Symmetry

As has been stated above, each of the paired bones was weighed
and measured separately. In the preceding tables the sum of the
weights of the two bones and the averages of the lengths have been
used. In table 4 we have listed the number of complete pairs of
bones in the second column. Where one bone was broken the weight
of the one normal bone was doubled and its length was used for
the data in tables 1-3, and so the numbers given in the second column
of table 4 are less than in the preceding tables. The percentage
frequency of a heavier or longer right bone, or left bone or bones
of the same weight or length, are given rather than the absolute
numbers, which can easily be obtained from the total number of
pairs of bones.

In no case do we find a percentage of 60 percent, and only three
times is there a percentage of over 50 percent in the weights, and
the highest percentage in the linear dimensions is 43.8 percent for
the right tibiofibula and for the femur. Four pairs of the bones
are heavier on the left side a little more frequently, but three pairs
of the six bones are longer a little more frequently on the left side.
The percentages in all cases are so low that it seems to be very
questionable whether there is any dominance of either side in either
weight or length of these six pairs of long bones.

Summary

The weights of the skull and of the six pairs of long bones have
the highest average coefficient of variation, followed by the average
of the twelve dimensions of the skull. The lengths of the long bones
are the least variable. The skull width is the most constant di-
mension of the skull.

A significant sex difference is indicated for but few of the weights
and linear measurements. Where there is a significant sex difference,
the females are more variable in both weight and length.

Only two of the correlations of the weights of the long bones and
the skull weight are significant. The correlations of the linear
dimensions show that the skull width, or the bitemporal diameter, is
the best criterion of the other linear dimensions. This is followed by
the skull height, and the lowest correlations are between the skull
length and the other linear dimensions. The low correlations for

196 The University Science Bulletin

both the weights and the linear dimensions would indicate that the
skull weight and the three linear dimensions of the skull are not good
criteria of the other weights and lengths.

There does not seem to be any significant asymmetry in the
weights or in the lengths of the paired bones.

LITERATURE CITED

Duckworth, W. L. H. 1917. Notes on some measurements made on subjects
in the dissecting room. J. Anat., vol. 51, pp. 167-179 and 376-391.

Dunn, H.albert L. 1929. Application of statistical methods in physiology.

Physiol. Revs., vol. 9, pp. 275-398.
Green, C. V. 1932. A genetic craniometric study of two species of mice and

their hybrids. J. Expr. Zool., vol. 63, pp. 533-551.

1933. Differential growth in the crania of mature mice. J. Mam-
malogy, vol. 14, pp. 122-131.

Latimer, H. B. 1937. Weights and linear dimensions of the skull and of some

of the long bones of the skunk {Mephitis mesomelas avia). J. Morph.,

vol. 60, pp. 379-391.
Latimer, H. B., and J. H. Rosenbaum. 1926. A quantitative study of the

anatomy of the turkey hen. Anat. Rec, vol. 34, pp. 15-23.
L.^TiMER, H. B., and Ray B. Riley. 1934. Measurements of the skull and of

some of the long bones of the muskrat (Ondatra zibethicus cinnamo minus) .

J. Morph., vol. 56, pp. 203-212.
Lerner, I. Michael. 1937. Relative growth and hereditary size limitation in

the domestic fowl. Hilgardia, vol. 10, pp. 511-560.
Orensteen, Meyer M. 1915. Correlation of anthropometrical measurements

in Cairoborn natives. Biometrika, vol. 11, pp. 67-81.
Orensteen, Meyer M. 1920. Correlation of ceiihalic measurements in Egyp-
tian-born natives. Biometrika, vol. 13, pp. 17-24.
Pe.\rson, Karl, and Adelaide G. Davin. 1924. On the biometric constants of

the human skull. Biometrika, vol. 16, pp. 328-363.
Schneider, M., and L. C. Dunn. 1924. On the length and variability of the

bones of the white Leghorn fowl. Anat. Rec, vol. 27, pp. 229-239.

Latimer and Asling: The Mourning Dove

197

-S.T-

^'^-B.S.— ^'^B.O.-^'

Fig. 1. Drawing of the dorsal side of the skull in upper part and lower side
in lower part, showing the ten dimensions measured.

EXPLANATION OF FIGURE

S. L., Skull length S.T., Spine sphenoid-tip of beak

H.T., Hinge-tip of beak B.S., Basion-spine of sphenoid

O.H., Occiput-hinge B.O., Basion-occiput

Bt., Bitemporal width F.L., Length foramen magnum

lo Intprorbitaj width F.W., Width foramen magnum

THE UNIVEESITY OP KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 10

Weights and Linear Dimensions of the Skull and of Some
of the Long Bones of the Red-tailed Hawk {Buteo
borealis borealis)

HOMER B. LATIMER,

Department of Anatomy, University of Kansas

Abstract: The skull and six long bones of twenty-four male and 27 female
hawk skeletons were weighed. Fourteen skull dimensions and twenty-one
dimensions of the long bones were made.

The weights and linear dimensions are as a rule more variable in the males.
The skull height is the most constant dimen-^ion in both sexes. Both skull
weight and skull length are fairly constant and their coefficients are less variable
than the average of the coefficients of variation.

The weights of the skull, the mandible and the six long bones are all
significantly heavier in the females. All of the thirty-seven linear dimensions
are greater in the females and most are significantly greater.

The weights and the linear dimensions are as a rule better correlated with
skull weight and skull length in the females. The skull weight is better
correlated with all of the weights than is the skull length, but the skull length
is better correlated with most of the linear dimensions. In the females, the
skull length is better correlated with the longitudinal dimensions of the skull
and the skull weight is better correlated with the transverse dimensions.

The frequency of heavier or longer members of the paired bones is given.
The asymmetries are not great enough to justify the conclusion that there
is anything but normal variation in a normal bilateral symmetry.

THE literature contains very few quantitative studies of the
avian skeleton, and so it has seemed best to utilize these skele-
tons which have been so well prepared, even though the numbers are
not as large as could be desired. This paper will give the average
measurements of the weights and of the linear dimensions of the
skull and of some of the long bones of the male and the female hawk
skeletons. Then the degree of dispersion of these measurements will
be given and this should show how reliable the various measurements
are for taxonomic studies. The sex differences will be presented

(199)

200 The University Science Bulletin

next. Next the correlations between the skull weight and the skull
length and the other measurements of the skull and of the long bones
will be presented. These correlations should give some idea of the
validity of using skull weight or skull length as general criteria for
the other skeletal measurements. Last of all the asymmetry of the
paired bones will be discussed very briefly. This follows the general
plan employed in the two earlier papers on mammalian skeletons,
or the muskrat (Latimer and Riley, '34), and the skunk (Latimer,
'37), and in the study of the skeleton of the mourning dove (Latimer
and Asling, '38).

This collection of hawk skeletons, like the collections used in the
preparation of the preceding papers, is from the Natural History
Museum of the University of Kansas. All of the bones of all four
species of animals have been prepared in the same manner. This
collection is not as large as the two mammalian collections, nor as
large as could be desired for final conclusions, but it is hoped that,
together with the preceding paper on the mourning dove skeleton,
it will contribute to the knowledge of the bird skeleton. Linsdale
('30) has made an excellent study of a series of 465 skeletons of
several subspecies of the fox sparrow. The skeletons of the road-
runner (Larson, '31) and of several species of woodpeckers (Burt,
'31) have been studied quantitatively, but this is the only study
of this kind, so far as is known, on the skeleton of any hawk.

Materials and Methods

The skeletons of fifty-one adult red-tailed hawks were studied.
Of these, twenty-four were males and twenty-seven females. A few
of the skeletons were injured in one or more bones or were slightly
incomplete, so the weights of the entire skeletons could not be
obtained. As many weights and measurements as possible were
made on each skeleton, and with the exception of the total body
length and the wingspread, the numbers of each measurement are
complete or are only a few short of the full number. The total body
length and the wingspread were recorded on the slip with each
skeleton and were made by the museum staff and were found in
only a little over half of the specimens.

All of the skeletons, packed in individual pasteboard boxes, were
kept in a dry room for some time before weighing, but the bones
were not oven dried. The weights were made on a chemical balance
sensitive to 0.1 milligram, but the weights were taken only to the
nearest milligram. In weighing and in measuring the paired bones,
the bone of the pair which was picked up first was weighed or

Latimer: The Red-tailed Hawk 201

measured first to avoid a definite sequence and a possible bias in
making the measurements. The weights of the paired bones in
tables 1 and 2 are the sum of the weights of the right and left
bones. The skull weight does not include the weight of the mandi-
ble.

All of the linear measurements were made with a vernier caliper
reading to 0.1 millimeter, and the dimensions of the paired bones in
tables 1 and 2 are the average of the right and left bones of each
pair. The methods used in making twelve of the dimensions of the
skull are illustrated in figure 1. The skull height was made with the
arm of the caliper parallel to and against the base of the skull with
the back of the caliper against the occiput, and the moveable arm
was carefully slipped down until it touched the top of the skull.
The length of the mandible was measured from the tip of the mandi-
ble to the most posterior part of the left side, with the back of the
caliper parallel to the left side of the mandible.

The lengths of the long bones were all made with the long axis of
the bone held parallel to the back of the caliper and the maximum
length taken. The widths of both the proximal and the distal ends
of the humerus, the femur and the tibiofibula are the maximum
widths and they were made with the bone held with its long axis
parallel to the arms of the caliper. The diameters of the shafts are
the minimum anteroposterior diameters. The diameter of the head
of the femur is the maximum anteroposterior diameter. The length
of the scapula is the maximum length and the width is likewise the
maximum width. The length of the sternum is the maximum length
in the midline, and the height was measured from the superior or
inner side of the bone to the free border of the keel, with the back
of the caliper held parallel to the anterior border of the keel. A
caliper with curved arms was used in making this measurement
which is largely the maximum height of the keel of the sternum plus
the thickness of the body of the sternum at the union of the body
and keel. The interacetabular diameter is the minimum diameter
from the dorsal sides of the rims of the two acetabula. The "ace-
tabulum-crest ilium" is the minimum distance from the rim of the
acetabulum to the crest of the ilium, and the width of the ischium
is the maximum diameter, including both right and left ischia.

The formulae used in making the computations are those usually
so employed and are given by Dunn ('29). All of the statistical
work was checked and all of the figures were carried to at least one
and usually two or three more decimal places than those given in
the tables.

202 The University Science Bulletin

I wish to express my most sincere appreciation to Mr. C. D.
Bunker for permitting the use of this excellently prepared collection
of skeletons, and to Mrs. Mary M. White for assistance in making
the linear measurements, and to Miss Edna Mae McConnell fox
making the weighings and for her careful help with the statistical
work.

Averages and Variability

The average measurements of both the weights and the lengths
and their coefficients of variation for both the males and the females
are given in table 1. These coefficients of variation of the hawk
skeleton have a higher average coefficient in the males, just as has
been found in the skunk skeleton (Latimer, '37), and in the external
dimensions of the cat (Latimer, '36). The average coefficient of
variation for the eight weights is 13.95 for the males and 13.42 for
the females, or a difference of 3.76 percent. The average of the
coefficients of the fourteen linear dimensions of the skull is like-
wise greater in the males, or 4.93 percent for the males and 4.84 per-
cent for the females, or the averages are 1.64 percent greater in the
males. The greatest difference is found in the last twenty-three
dimensions in this table with an average coefficient of variation of
5.37 for the males and an average of 5.11 for the females, or a differ-
ence of 6.63 percent. As is to be expected, the weights are more
variable than the linear dimensions. The dimensions of the skull
are in general less variable than are the other linear measurements,
and of the skull dimensions the skull height is the most constant in
both sexes. The skull length is less variable than any of the last
twenty-three dimensions in both sexes, and of the skull dimensions
in the male only four are less variable than the skull length and but
three in the female. This would show that the skull height and, to
a lesser degree, the skull length, are constant and fairly reliable
dimensions.

The longitudinal measurements of the skull and of the long bones
are less variable on the average than are the transverse dimensions.
The length from the basion to the occiput (B.O. in figure 1) does
not seem to be a constant measurement in the hawk or in the mourn-
ing dove, for it was the most variable of the skull dimensions in the
dove and also the most variable in the male hawk and second most
variable in the female hawk. The interorbital and the binasal di-
ameters are also variable. They are the two most variable dimen-
sions in the female and the second and third in rate of variability in
the male hawk. Both of these measurements are made from the

Latimer: The Red-tailed Hawk 203

thin edges of the bone. The central portions of these bones provide
adequate strength and the thin edges may vary without making any
significant difference. These are rather unreHable measurements.
The edges of the bone bordering the orbits, especially, show irregu-
larities which would make this dimension variable. The two diam-
eters of the foramen magnum in the hawk skull are fairly constant.

The greater variability in the transverse dimensions of the long
bones may, in a measure, be accounted for by the greater difficulty
in making these measurements and because of their smaller size.
This would not explain the variability of the width of the skull.
In the females, the proximal bone of each appendage is less variable
in weight and in length, with the exception of the ulna, than the
more distal bones in the same extremity. This is not true for the
males, however. The ulna is the most constant in weight of all the
bones in the males and the humerus in the females. The three wing
bones are less variable in the males than the three bones of the lower
extremity, and this is also true for the females if the average vari-
ability of the three bones in each extremity is considered. The man-
dible is the most variable in weight in the males and in the females
the tibiofibula is the most variable in weight. The measurements of
the total length and the wingspread have high coefficients of vari-
ation.

A comparison of these coefficients of variation with similar coeffi-
cients of the other skeletons studied shows that the variability of
these measurements in the hawk is about the same as that found in
the two mammalian skeletons and the one bird skeleton. The hawk
skull is about as variable as that of the fox sparrow, but the lengths
of the long bones seem to be more variable in the hawk than in the
fox sparrow (Linsdale, '30).

The last column of table 1 gives the significant ratios, or the
difference between the average measurements of the female and the
male divided by the probable error of this difference. The greatest
differences are in the weights and all of these are significantly heavier
in the females. The greatest sex differences seem to be in the weights
of the three wing bones. Of the next fourteen linear dimensions
of the skull five, or 35.71 percent, are not significantly greater in
the females. Table 1 shows the basion-occiput length the same in
both sexes, but in computing these ratios the figures used were the
figures before they were rounded out and cut to but one decimal
place. The average lengths of the basion-occiput with three decimal
places are 9.207 for the female and 9.190 for the male, thus making
this dimension slightly greater in the female and giving a ratio of

204 The University Science Bulletin

0.10, as shown in the last column of this table. Only three of the last
twenty-three dimensions do not show a significant sex difference,
and one of these, namely the scapula width, is nearly great enough
to conclude that there is very probably a real sex difference in the
width of the scapula in the two sexes.

All of the weights and the linear dimensions of the male and the
female hawks were plotted as frequency polygons (not shown for
lack of room). Fifty-six percent of these polygons in each sex were
skewed toward the smaller measurements. The remainder were
skewed chiefly toward the greater measurements. A very few were
symmetrical.

In general it may be said that the dimensions of these bones are
fairly constant as compared with other animal skeletons. The
greater variability is found in the male skeletons. The skull weight,
length and height are all more constant than the average of the other
coefficients for each group. The skull height is the most constant
dimension of the skull in both sexes, and is also more constant than
any of the other linear dimensions. All of the weights are signifi-
cantly heavier in the females, and all of the linear dimensions are
greater in the females and the majority of these are significantly
greater.

Correlations

The correlations of the eight weights and the thirty-seven linear
dimensions with skull weight and with skull length for both males
and females are given in table 2. It has been shown in table 1 that
the weights and the linear dimensions are in general more variable
in the male hawks, and this table shows that the weights, the linear
dimensions of the skull and the other group of linear dimensions
all have higher average correlations with both skull weight and skull
length in the females. The only exceptions in the correlations of
the weights are the weight of the radius with skull length and the
weight of the femur with skull weight. There are 'several exceptions
in the linear dimensions, and most of these are correlations with
skull length. In general the ma'les are more variable and the skull
weight and the skull length are not as well correlated with the other
weights and linear dimensions as in the female hawks.

The averages of the correlations of the eight ponderal measure-
ments show that they are better correlated with skull weight than
with skull length in both sexes. The last twenty-three dimensions
in table 2 have a higher average correlation with the skull length in
both the males and the females. In the male hawks, the skull length

Latimer: The Red-tailed Hawk 205

has a higher a^T'ragc correlation witli th.e fourteen linear dimensions
of the skull, but in the females the averages of the correlations are
practically the same with both skull weight and skull length.

The last group of twenty-one linear measurements, omitting the
last two, which are not skeletal dimensions, consists of two different
types of measurements, namely, those parallel to the long axes of the
bones and those perpendicular to the long axes, or measurements of
length and measurements of the thickness or of the strength of the
bones. The question arose as to whether the skull weight would be
found to be better correlated with the transverse diameters and the
skull length with the longitudinal dimensions. The averages of these
two types of dimensions show that in the males the skull length has
a higher average correlation with both the longitudinal and the
transverse measurements of the long bones and also higher average
correlations with both the longitudinal and the transverse dimensions
of the skull. Hence in the male hawk the skull length is in general
a better criterion of all of the linear dimensions than is the skull
weight. In the female hawks the skull length is better correlated
with the seven linear dimensions of the skull and the skull weight is
better correlated with the six widths of the skull. The averages of
the correlations of the ten longitudinal measurements and the eleven
transverse dimensions in the last twenty-one dimensions of this
table, with both skull weight and skull length in the females, show
little difference. Thus the skull weight and the skull length are
nearly equally good indices of both the longitudinal and the trans-
verse dimensions in the female hawks.

All of the correlations in the female skeletons are positive, but
some are so low that they are not significant. All but one of the
male weights are positive when correlated with either skull weight
or skull length. Six of the linear correlations in the males are nega-
tive, three of these are correlations with skull weight and three dif-
ferent dimensions are correlated with skull length. None of these
negative correlations are significant and all but one have larger prob-
able errors than the correlation itself. The highest of the correla-
tions with the weights is -f- 0.874 for the female skull weight cor-
related with the weight of the humerus. The highest correlation in
this table is + 0.929 and it is found between the male skull length
and the length from the end of the palatine bone to the tip of the
beak. The highest correlation in the female is this same dimension
correlated with skull length. Although this dimension (P.T. in figure
1) forms a relatively large proportion of the total skull length, the
point used as the posterior end of this measurement must be fairly

206 The University Science Bulletin

constant and also the length of the beak. In the mourning dove skull
(Latimer and Asling, '38) the beak was found to be quite variable,
but this measurement in the hawk, which includes the length of the
beak, has a coefficient of variation of but 3.86 in the males and
3.91 in the females (table 1).

If the averages of the correlations for the various groups of cor-
relations in table 2 are compared with similar average correlations
given in the other two papers on the mammalian skeletons, these
averages are lower, indicating that these measurements of the skull
in the hawk are not as good criteria of the other measurements as
are the similar skull dimensions in the muskrat and in the skunk, but
they are much better than the similar averages for the dove skeleton.
The skull lengths in the male hawks have the lowest averages of
any of the groups, or -j- 0.241 when correlated with skull weight and
-J- 0.478 when correlated with skull length. The next lowest are the
other linear dimensions in the males (last 23 correlations in table 2),
which when correlated with skull weight have an average correlation
of -\- 0.425 and a correlation of -|- 0.487 when correlated with skull
length. All of the other groups of linear dimensions have average
correlations above + 0.5 in both the males and the females. The
weights in the males when correlated with skull weight have an
average correlation of -\- 0.595, and -f- 0.446 when correlated with
skull length. Similar correlations in the females are, respectively,
-|- 0.741 and -)- 0.587. These correlations would show that in the
two avian forms studied the skull weight or the linear dimensions of
the skull are not as good criteria of the weights and the linear di-
mensions of the other parts of the skull and of the other bones of
the skeleton as they are in the two mammals studied.

In general, the correlations with both skull weight and skull length
are higher in the females. The skull weight gives higher correla-
tions with the other weights and in the males the skull length is a
better criterion of all of the linear dimensions, both longitudinal
and transverse, than is the skull weight. In the females, the skull
length is better correlated with the other seven longitudinal dimen-
sions of the skull and the skull weight is better correlated with the
six transverse dimensions.

Asymmetry

The right and the left bones of each pair were weighed and
measured separately as described above and the sum of the weights
or the average of the lengths of each pair given in tables 1 and 2.
In table 3 the number of pairs of bones is given in the second column

Latimer: The Red-tailed Hawk 207

and the following columns give the percentage frequency of a
heavier or longer right bone, a heavier or longer left bone, or bones
of equal weight or of equal length. This table shows a greater
asymmetry in the bones of the hawk than is shown for the mourning
dove skeleton (Latimer and Asling, '38). The humerus and radius
are both heavier more frequently on the right side and the three
bones of the lower extremity are heavier more frequently on the left
side, suggesting a crossed symmetry (Schaeffer, '28) for a bird which
does not use the two pairs of extremities at the same time or for the
same method of locomotion. The second part of this table shows
that the lengths of the paired bones of both the wing and the leg are
longer more frequently on the right side, the lengths of the scapula
and the coracoid alone being greater more frequentty on the left side.
In general, the hawk shows a greater asymmetry in both weight
and in length of the long bones of the extremities than does the
mourning dove. In degree of asymmetry it resembles the three
mammals studied, and like the other skeletons there seems to be no
significant type of asymmetry, but probably merely a fortuitous
variation. These variations are frequently so small that they would
not be functionally significant. According to Griineberg ('35) these
asymmetries are a common type and they are "caused directly
through the asymmetrical interference of environmental factors
with the action of equally distributed genes on a symmetrical sub-
stratum."

Summary

The weights and the linear dimensions are as a rule more variable
in the males. The skull height is the most constant linear dimension
in both sexes. Both skull weight and skull length are fairly con-
stant and the coefficients of both are less variable than the average
of the coefficients of variation.

The weights of the skull, mandible and the six long bones are all
significantly heavier in the female hawks. All of the thirty-seven
linear dimensions are greater in the females and most of these are
significantly greater.

The weights and the linear dimensions are as a rule better cor-
related with skull weight and with skull length in the female hawks.
The skull weight is better correlated with all of the weights than is
the skull length, but the skull length is better correlated with most
of the linear dimensions. In the females, the skull length is better
correlated with the longitudinal dimensions of the skull and the
skull weight is better correlated with the transverse dimensions.

208 The University Science Bulletin

The frequency of a heavier or a longer member of the paired
bones is given. A crossed symmetry seems to be apparent for the
weights of the humerus and the femur, but the lengths of all of the
six long bones of the two extremities are greater more frequently on
the right side. None of the percentages are probably great enough
to warrant the conclusion that there is anything but normal varia-
tions in a normal bilateral symmetry.

LITERATURE CITED

Bltrt, Willlam Henry. 1931. Adaptive modifications in the woodpeckers.
Univ. of Calif. Pub. in Zool., vol. 32, pp. 455-524.

Dunn. H. L. 1929. Application of statistical methods in physiology. Physiol.
Reviews, vol. 9, pp. 275-397.

Grlneberg, Hans. 1935. The causes of asvmmetrics in animals. Am. Nat.,

vol. 69, pp. 323-343.
Larson, Leigh Marian. 1931. Osteologv of the California road-runner recent

and Pleistocene. Univ. of Calif. Pub. "in Zool., vol. 32, pp. 409-428.
L.ATiMER, H. B. 1936. Weights and linear measurements of the adult cat.

Am. J. Anat., vol. 58, pp. 329-347.

1937. Weights and linear dimensions of the skull and of some of the

long bones of the skunk {Mephitis mesonielas avia) J. Morph., vol. 60, pp.

379-391.
L.^TiMER, H. B., and O. W. Asling. 1938. Weights and linear dimensions of

the skull and of some of the long bones of the mourning dove {Zenaidura

macroura carolinensis) . Bull. Univ. of Kan., vol. 25, pp. 187-197.
Latimer, H. B., and R. B. Riley. 1934. Measurements of the skull and of some

of the long bones of the muskrat (Ondatra zibcthicus cinnamo minus) . J.

Morph., vol. 56, pp. 203-212.
LiNSDALE, Jean M. 1930. Variations in the fox spaiTOw (Passerella iliaca)

with reference to natural historv and osteologv. Univ. of Calif. Pub. in

Zool., vol. 30, pp. 251-392.
Schaeffer, a. .a. 1928. Spiral movement in man. J. Morph. and Physiol.,

vol. 45, pp. 293-398.

Latimer: The Red-tailed Hawk

209

Table 1
Average measurements, coefficients of variation and significant ratios

Skull

Mandible

Humerus

Radius

Ulna

Femur

Tibiofibula

Tarsometatarsus

Males.

Average weight
in grams.

5.195
1.049

635
307
6.556
5 353
8 017
5.627

Average length
in millimeters.

Skull length

Skull height

Basion-occiput

Basion-palatine

Palatine-tip of beak

Occiput-nasals .

Mandible, length

Bitemporal width

Interorbital width

Bilacrymal width

Binasal width

Biauditory width

Foramen magnum Icnpth
Foramen magnum width

0.110
0 027

0 130
0 0.36
0 108
0 154
0.175
0.119

78 3
31 8

9 2
19 5
41.6
55 5
61 9
49.1
17.9
45 9

5 0
41 6

7 3

0 37
0 10
0 15
0 17
0 30
0 24
0 33
0.18
0 26
0 43
0 06
0 17
0 05
0 05

CoefiBcient
of variation.

13.31
17.58

11 61

10 67
10 60
18.62
14 83
14.36

1 52
1 93

1 25

1 15
1,17

2 11
1.62
1.56

3 21

2 02
10 37

5 49

3 86

2 f8

3 59
2 52
9 79
5 31
8 34

2 85

4 PC

3 82

0 34

0 22

1 15
0 62
0 42
0 32

0 38
0.27

1 C5
0 66

0 to

0 30
0 53
0 41

Femsles.

.^.verage weight
in grams.

6.070
1.250

0 088
0 024

9.518 ± 0 127
3 099 ± 0 052
9 201 ± 0 173

6 551 =t 0 104
10.139 ± 0.217

7 331 ± 0 145

Average length
in millimeters.

51 0

18

47 0
5 8

43 7
7 3

3 ±

0 36
0 10
0 08
0 15
0 26
0 18
0 36
0 19
0 20
0 35
0 08
0 20
0 04
0 05

Coefficient
of variation.

12.17
14.29

10 30
12 72
14.47
12 05
16.15
15.24

1 15
1.36

0 96

1 21
1 33
1 14
1 55
1 43

40

34

60

55

91

41

32

76

8 28

5 15

10.97

3 47

4 71
3 f4

0 31
0 21
0 61
0 51
0 36
0 22
0 40
0.26
0.77

0 53

1 02
0 33
0 43
0 36

Difference

P. E. diff.

5 94
5 57

10 36

12.52

12.97

6.45

7 61

9 08

4 85

3 52

0 10
2.52
4.r4
3.86

4 75
7 55

36
.87
36
71
FO

3.16

Humerus, length

Humerus, width proximal end . .
Humerus, anteroposterior diam.

Humerus, width distal end

Radius, length

Ulna, length

Femur, length

Femur, diameter of head

Femur, anteroposterior diam. . .

Femur, width distal end

Tibiofibula, length

Tibiofibula, width distal end. . . .

Tarsometatarsus, length

Scapula, length

Scapula, width

Coracoid length

Sternal length

Sternal height

Interacetabular diameter

.fVcetabulum-crest of ilium

Ischium, width

Total length.
Wincspread .

106 9

24 0

7 1

18 7

1)1 5

124 8

82.

6

7

17
109
13
82
60

8
45
68
19

26 0
40.9
39.3

.".'>2 2
376.8

0 65
0 13
0 05

0 12

1 09
0 79
0 50
0 06
0 07
0 15
0 62
0 10
0 52
0 38
0 07
0 41
0 61
0 19
0 30
0 29
0.30

6 86
5 55

4 13

3 81

4 66
4 45
6 65

10

fO
81
58
89
83
82
28
35
61
06
03
70
95

4.82
5.14

7.59
7,24

0 44
0 41
0 50
0 48
0 71
0 45
0 43
0 62
0 61
0 63
0 41
0 52
0 46
0 47
0 60
0 63
0 65
0 72
0 85
0 52
0.55

0.91
1.05

113 5

25 7

7.6

20 0

124 5

131 8

85.9

7 2

8 4
18.7

115 3
14
84
63
9
48
73
20
26

43.8
41.5

562.3
411.3

0 51
0 15
0 05
0 12
0 59
0 59
0 41
0 06
0 07
0.14
0 57
0 12
0 45
0 37
0 OS
0 29
0 51
0 15
0 16
0.23
0.31

3.13
9.29

49
62
06
49
64

3 47

62
93
15
57
71
48
12
46
80
62
34
52
53
98
76

4.20
12.08

0 31
0 43
0 47
0 41
0 33
0 32
0 34
0 55
0 57
0 .M
0 35
0 60
0 38
0 41
0 63
0 J2
0 49
0 51
0 42
0.37
0 53

0.39
1.62

7.94
8 35
6 33
7.88
10 45
7.07
5 86

5 95
8.51
8 10

6 55

7 79
3 80
6 25
2 ro
6 08
5.50
5 34
2 27

8 04
4.98

1.34
3.19

14—4141

210

The University Science Bulletin

Table 2
Correlations

Males.

Females.

Weights.
(All weights in grams).

Correlations
with skull weight.

Correlations
with skull length.

Correlations
with skull weight.

Correlations
with skull length.

Skull

4-0 307 ± 0 133
4-0 639 ± 0 083
4-0 665 ± 0 078
4-0 398 ± 0 118
— 0 118 =t 0 142
4-0 467 ± 0 112
4-0 526 ± 0 102
4-0 681 ± 0 075

4- 0 616 ± 0 082

4-0 547 =t 0 103
4- 0 676 ± 0 080
4-0 367 ± 0 127
4-0 575 ± 0 101
4-0 594 ± 0 098
4-0 711 ± 0 073
4- 0.697 ± 0.076

4-0 812 ± 0 045
4-0 874 ± 0 031
4- 0 808 ± 0 048
4-0 419 =t 0 109
4-0 812 ± 0 047
4-0 757 ± 0 059
-hO 707 ± 0 066

4- 0 660 ± 0 075

4- 0 639 ± 0 077

Radius

4- 0 533 =fc 0 095

Ulna

4- 0 336 ± 0 115

Femur

4- 0 635 ± 0 079

Tibiofibula

4- 0 638 ± 0 078

4- 0 638 ± 0 077

Linear Dimensions.
(All lengths in millimeters).

SkulUength

4-0 307 =fc 0 133
4-0 431 ± 0 120

— 0 086 ± 0 146

— 0 221 ± 0 140
4-0 534 =t 0 105
4-0 170 ± 0 143
4-0 724 =t 0 070
4-0 254 ± 0 138
4- 0 280 =fc 0 136

— 0 090 ± 0 167
4-0 329 =fc 0 131
4- 0 373 =t 0 127
-i- 0 323 ± 0 132
4-0 045 ± 0.147

4-0 616 ± 0 082
4- 0 700 ± 0 067
+ 0 094 =t 0 131
4-0 100 ± 0 131
+ 0 746 ± 0 059
4- 0 626 ± 0 080
4- 0 842 ± 0 0.39
4-0 812 ± 0 047
4- 0 640 =fc 0 078
4- 0 568 ± 0 097
4- 0 590 ± 0 086
4-0 573 ± 0 092
4-0.237 ± 0 125
4-0.618 =t 0 082

Skul! height

4- 0 400 ± 0 118
4- 0 572 ± 0 097
-h 0 643 ± 0 086
4- 0 929 =t 0 020
4- 0 704 ± 0 071
4- 0 823 =t 0 045
4- 0 048 =t 0 140
4- 0 278 =t 0 130
— 0 153 ± 0 160
4- 0 634 ± 0 084
4- 0 759 =fc 0 060
+ 0 173 =t 0 140
4-0 405 ± 0.118

4- 0 548 ± 0 091

Basion — occinut ....

4- 0 229 =t 0 123

Basion — palatine

4- 0 725 ± 0 062

Palatine — tip of beak

4- 0 907 =t 0 023

Occiput — nasals

4- 0 615 ± 0 081

Mandible, length

4- 0 785 ± 0 050

4- 0 682 ± 0 071

4- 0 430 =fc 0 106

Bilacrvmal width

J- 0 481 =fc 0 111

Binasal width

4- 0 480 ± 0 100

4- 0 743 ± 0 059

4- 0 002 ± 0 130

4- 0 515 ± 0 095

Humerus, length

Humerus, width proximal end

Humerus, anteroposterior diam. shaft

Humerus, width distal end

Radius, length

Ulna, length

Femur, length

Femur, diameter of head

Femur, anterppopterior diam. shaft.

Femur, width diptal end

Tibiofibula length

Tibiofibula, width distal end

Tarsometatarsus, length

Scapula, length

Sea pula, width

Coracoid, length

Sternum, length

Sternum, height

Inter.icptabu'ar diameter

Acetabulum — crest ilium

Ischium, width

Totallength

Wingspread

4- 0 285
4-0 623
4- 0 743
4- 0 608
4- 0 152
4-0 301
4- 0 348
4- 0 554
4- 0 686
4- 0 408
4- 0 292
+ 0 454
4-0 384
4- 0 296
4-0 243
+ 0 365
4-0 567
4-0 478
4- 0 379
4- 0 413
4-0 348

0 135

: 0 OPO

^ 0 066

0 095

0 144

0 137

0 133

: 0 102

: 0 080

. 0 123

0 135

: 0 117

0 125

0 134

0 1.38

: 0 128

: 0 100

: 0 114

: 0 126

0 122
: 0 129

4-0 517 =t 0 123
4-0 320 ± 0 202

4-0 610 :
4-0.209 ■■
4- 0 662
4- 0 708 ^
4- 0 343 ^
4- 0 586
4- 0 523 :
4- 0 464
4- 0 817
4- 0 570 ^
4- 0 637 :
4- 0 573
4- 0 679
4- 0 521 ^
4- 0.376 ^
4- 0 493
4- 0 630 ■■
+ 0 .507 ■

— 0 022 ^
4- 0 470
4- 0 687 -

— 0 100 ^
4- 0 266 ^

0 088
0 135
0 079
0 070
0 124
0 094
0 104
0 110
0 048
0 095
0 084
0 Or4
0 076
0 102
0 121
0 109
0 085
0 104
0 141
0 110
0 074

0 157
0 189

4- 0 661 ^

4- 0 692 :

4- 0 489 :

-1- 0 666 :

4-0 678 ^

4- 0 642 ^

4- 0 784 ^

4- 0 515

4- 0 .^04

4- 0 705

4- 0 664 ^

4- 0 552 ■■

4- 0 589 -

4- 0 657 ■■

4- 0 804 :

4- 0 660 :

4- 0 697 ^

4-0 511 :

4- 0 410 ^

4- 0 782 :

4- 0 278 :

0 074
0 069
0 101
0 074
0 072
0 078
0 053
0 097

: 0 099
0 067
0 075
0 092
0 086
0 075
0 047

: 0 075
0 068
0 098

0 no

0 051
0 122

4- 0 204 =t 0 132
4- 0 116 ± 0 192

4-0 640
4- 0 461
4- 0 505
4- 0 640
4- 0 639 ■
4-0 616
4- 0 714
+ 0 640
+ 0 576
4- 0 590
4- 0 636
4- 0 ,"^47
4- 0 691 ^
4- 0 728 ^
4-0 546
4-0 723
4-0 744
-I- 0 540
4- 0 6P5
-4-0 678
4-0 372

4- 0 595
4-0 350

0 077
0 102
0 097
0 077
0 077
0 081
0 065
0 077
0 087
0 085
0 079
0 091
0 068
0 061
0 091
0 062
0 058
0 092
0 067
0 070
0 112

0 085
0.164

Latimer: The Red-tailed Hawk

211

Table 3
Asymmetry in weight and length of the long bones

The heavier or longer right or left bone, or bones of equal size, are given in terms of per-
centage of the total number of measurements shown in second column.

Weights.

Humerus

Radius

Ulna

Femur

Tibiofibula

Tarsometatarsus

Linear Measurements.

Humerus, total length

Humerus, width, proximal end

Humerus, width, distal end

Humerus, anterioposterior diameter. .

Radius, total length

Ulna, total length

Femur, total length

Femur, diameter of head

Femur, width, distal end

Femur, diameter of shaft

Tibiofibula, total length

Tibiofibvila, width, distal end

Tarsometatarsus, total length

Scapula, length

Scapula, width

Coracoid, length

Number

Right

Left

of

greater,

greater.

Same.

cases.

percent.

percent.

50

64.00

36.00

0

41

51.22

48.78

0

40

40.00

60.00

0

47

46.81

53.19

0

46

45.65

54.35

0

50

46.00

54.00

0

50

44.00

38.00

18.00

51

50.98

29.41

19.61

50

36.00

42.00

22.00

50

24.00

28.00

48.00

41

60.98

26.83

12.19

40

72.50

17.50

10.00

47

48.94

23.40

27.66

50

22.00

38.00

40.00

51

50.98

29.41

19.61

49

30.62

32.05

36.73

46

56.52

28.26

15.22

50

34.00

38.00

28.00

50

62.00

18.00

20.00

44

31.82

59 . 09

9.09

50

58.00

22.00

20.00

47

27.66

51.06

21.28

212

The University Science Bulletin

•bt.W

Fig. 1. Drawings of the dorsal and ventral aspects of the hawk skull to
show the methods of making the twelve measurements. Two additional meas-
urements were made and thej^ are described in the text.

Description of the Figure

S.L., Skull length lo.W., Interorbital width

B. O., Basion-occiput L.W., Lacrymal width

B.P., Basion-palatine B.N., Binasal width

P.T., Palatine-tip of beak A. W., Auditory width

O.N., Occiput-nasals F.M.W., Foramen magnun width

Bt.W., Bitemporal width F.M.L., Foramen magnum length

THE UNIVEESITY OP KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 11

On the Alkyl Derivatives of the Isomeric Ortho and
Para-phenoxyphenyl Thiazolidones

MERRITT ELISHA ROBERTS and F. B. DAINS,
Contribution from the Chemical Laboratory of the University of Kansas

PREVIOUS investigations from this laboratory (1) have shown
that the 2-arylamino thiazoHdones

S — C(NHR): N — CO — CH2

react with alkyl iodides yielding in varying amounts ; (A) the 2-
aryl-2-alkyl thiazolidone and (B) the 2-arylimino-3-alkyl thia-
zolidone.

When the aryl group contained halogen or nitro substituents, there
seemed to be a tendency toward the formation of type B. For the
present investigation a study has been made of the effect of a
phenoxy grouping in both the ortho and para positions, in order to
ascertain the influence of this relatively high molar weight radical.

The stable thiazolidones,

S — C (NHC,H,OCoH,) : N — CO — CH,

were prepared by the two following methods: AVhen the mono-
phenoxyphenyl thioureas (RNHCSNHo) from the hydrochloride of
phenoxyaniline and ammonium thiocyanate were heated in alcohol
solution with ethylchloroacetate, the stable form alone was isolated.
A second preparation was as follows : Phenoxyaniline and chloro-
acetylchloride in acetone solution gave the omega-chloroacetphe-
noxyanilide, ClCH.CONHCeH.OCeH^. When this was heated in
ninety-five percent alcohol with potassium thiocyanate, ring closure
occurred with the formation again of the stable thiazolidone.
Modifying the conditions of this latter case, which will be described
later, led to the isolation of the 2-imino or labile form,

S — C (NH j : NR — CO — CH,.

(213)

214 The University Science Bulletin

The sodium salt of the 2-p-phenoxyphenylamino-4-thiazolidone
on alkylation gave a mixture of the 2-aryl-2-alkyIamino and 2-
arylimino-3-aIkyl thiazolidones. The ratio of the two isomers
formed was twenty-five parts of 2-alkyl to one part of the 3-alkyl
derivatives. From the sodivun salt of o-phenoxyphenyl-4-thiazo-
lidone the 2-aryl-2-alkyl isomer alone was isolated.

Alkaline hydrolysis caused the ring to break unless it was sub-
stituted at position No. 5. Acid hydrolysis left the ring intact,
either when substituted or unsubstituted, but hydrolyzed off the
phenoxyaniline group at position No. 2.

Methylene reactions could be obtained at position No. 5 using
benzaldehydes. Substituted benzaldehydes reacted with greater
ease but were apparently less stable.

Labile Thiazolidones: An especially interesting part of the re-
search had to do with the labile thiazolidones,

S — C(NH) : N(CeH,OCeHJ — CO — CH„

which were made by the action of potassium thiocyanate on the
omega-chloracet derivatives in absolute alcohol. The ortho-phe-
noxyaniline derivatives proved to be much more stable than the
corresponding para derivatives. The para readily rearranged to
the stable type by heating in dilute alcohol or by allowing it to
stand for several days in ninety-five percent alcohol. The ortho
rearranged best by use of acetic acid. The unstable type be-
came relatively stable as soon as the imino group was substituted.
The imino group reacted with RNCO and RCOCl compounds very
readily, forming urea and acylimino derivatives, respectively. Also
under certain conditions substitution at position No. 5 could be
made without rearrangement. Substitution at position No. 5 sta-
bilized the ring so that it was not broken by acid hydrolysis. All
methods of hydrolysis for the unsubstituted, labile types failed to
yield evidence as to structure, except in the case of acid hydrolysis,
which yielded substituted hydantoic acids. Substitution at position
No. 5 raised the melting point of the compounds, whereas substitu-
tion at position No. 3 lowered the melting point. These unstable
compounds gave non-isolated sodium salts as shown by their re-
actions. A stable HCl salt could be formed by using hydrogen
chloride gas, which could be neutralized to give the original unstable
form.

Roberts and Dains: Isomeric Ortho 215

EXPERIMENTAL
Preparation of Intermediates

p-Phenoxynitrobenzene, CeHgOCgH^NO, (I)

This was made by the Ulhiian method by heating a mixture of
potassium phenolate, copper powder and p-chloronitro benzene. On
reduction with zinc dust and calcium chloride (2) it gave p-phenoxy-
aniline (II) which boiled at 187-189° at 14 m.m. and melted at
83.5° C. Some derivatives of this were made for identification
purposes.

a-p-Phenoxyphenyl-/3-Phenylurea,

CeH.OCeH.NHCONHC.Hg (III)

This was made by the condensation of phenyl isocyanate and the
phenoxyaniline in dry benzene solution. The white crystals from
alcohol melted at 194°.

Anal. Calcd. for C,gH,,K,0,: N, 9.21. Found: 9.12, 9.23.

p-Phcnoxyphenyl-ethylamine, CeH^OCeH^NHC^H^ (IV)
This was the product formed by refluxing p-phenoxyaniline with
ethyl iodide. The yellow oil boiled at 213-214° at 32 m.m.

a-p-Phenoxvphenyl-a-ethvl-^-Phenylurea,

C,H,OC,H,N — CONHCgH, (V)
When the secondary amine (IV) was treated with phenylisocya-
nate, a little of the urea melting at 194° (III) was obtained, show-
ing the presence of traces of the phenoxyaniline. However, the
main product was the more soluble urea (V), which melted at 118°.
Anal. Calcd. for C^iH^iNoOo: N, 8.43. Found: 8.24, 8.43.

p-Phenoxyphenylisothiocyanate, CeHgOCgH^N C S (VI)
Efforts to prepare this by heating the di-p-phenoxyphenylthiourea
(m.p. 163-164°) with sulfuric acid or acetic anhydride were fruit-
less. A very poor yield resulted from the interaction of thiophosgene
with the phenoxyaniline, but fairly good results (60%) were ob-
tained by the action of lead nitrate on the ammonium di-thio-p-
phenoxyphenyl carbamate (3). The mustard oil was extracted from
the lead sulfide with hot alcohol. It had a characteristic odor and
melted at 42°.

Anal. Calcd. for C^oHgONS: N, 6.15. Found: 6.22, 6.32.

a-/>-Phenoxyphenyl-^-ethyl thiourea,

CeHsOCsH.NHCSNHC^Hg (VII)

216 The University Science Bulletin

This was easily prepared by adding an excess of ethylamine to
an alcoholic solution of the above mustard oil. The compact crys-
tals melted at 102°.

Anal. Calcd. for Ci^Hi.N.OS: N, 10.30. Found: 10.10,10.24.

a-p-Phenoxyphenyl-/?-benzyl thiourea,

CeH.OCsH.NHCSNHCH^CsH, (VIII)

The thiourea formed by the interaction of the isothiocyanate and
benzylamine in ether-benzene solution melted at 150° when recrys-
talized from benzene.

Anal. Calcd. for C.oHj.N.OS: N, 8.40. Found: 8.23, 8.21.

a-p-Phenoxypheny 1-a-benzy 1-^-phenyl urea ,

CgH^OCeH.C.H.NCONHC.H, (IX)

When paraphenoxyaniline was refluxed in alcohol solution with
benzyl chloride, phenoxyphenylbenzylamine was obtained as an oil
boihng at 293-305° at 40 m.m. This was dissolved in dry benzene
and treated with phenyl isocyanate. The resulting urea melted
at 185°.

Anal, calcd. for C,,H,2N,0o; N, 7.10. Found: 6.82.

Thiazolidone Synthesis (Stable)

?>Phenoxychloroacetanilide, QHgOCeH.NHCOCH.Cl (X)
The paraphenoxyaniline (1 mol.) was dissolved in a mixture of
acetone and pyridine (2 mols.). .To this was slowly added chloro-
acetylchloride (iVs mols.). After standing for one hour the crude
product was purified by crystallization from alcohol. The fine white
needles melted at 101°.

Anal. Calcd. for Ci.Hi^NOoCl: N, 5.36. Found: 5.30, 5.43.

2-23-Phenoxyphenylamino-4-thiazolidone,

S — C(NHC,H,OC,H,)NCOCtL (XI)

This is the stable form, and was made by refluxing for four hours
the chloroacet-derivative (X) and potassium thiocyanate in alcohol
solution. On pouring into water the yellow thiazolidone separated,
which melted at 183.5° after crystallization from alcohol. The same
thiazolidone was also prepared by the action of ethylchloroacetate
on monophenoxyphenylthiourea.

Anal. Calcd. for C15H12N AS: N, 9.86. Found: 9.75,9.77.

The labile or 2-imino-3-paraphenoxyphenyl-4-thiazolidone will
be discussed later.

I

Roberts and Dains: Isomeric Ortho 217

Acid Hydrolysis. The thiazolidone (XI) was hydrolyzed by the
action of hydrogen chloride in hot acetic acid into p-phenoxyaniline
and 2, 4-thiazoldione (m.p, 125°).

Benzal Derivative. It condensed readily with benzaldehyde in
alcohol solution with a few drops of sodium hydroxide, forming
2-y0-phenoxyphenylaniino-5-benzal-4-thiazolidone,

' S — C(NHC,H,OaH,)NCOC = CHCeH,

I 1

(XII) which melted at 241°.
Anal. Calcd. for Co.oH.^N.O.S; N, 7.53. Found: 7.73, 7.75.

Sodium Salt. This was obtained by dissolving the thiazolidone
(XI) in hot twenty-percent sodium hydroxide. On cooling, the
sodium salt separated, and could be purified by crystallizing from
a mixture of benzene and alcohol.

Ethyl Derivatives. The dry sodium salt was dissolved in ethyl
alcohol and an excess of ethyl iodide added. After refluxing eight
hours it was steam distilled and the residue taken up in ether. The
ether solution was extracted with acid and the extract neutralized
with ammonium hydroxide. The compound which separated melted
at 108° (yield 28 grs.) and was proven to be 2-23-phenoxyphenyl-2-
ethylamino-4-thiazolidone,

S_C(NaH,0C,Hg,C2H,) NCOCH3 (XIII)

Anal. Calcd. for C,,H,,N,0,S: N, 8.95. Found: 8.94, 8.91.

From the residual ether solution was isolated one gram of the
3-ethyl derivative (XIV), melting at 124°.

Proof of Constitution of XIII. Since acid hydrolysis failed to
give satisfactory results, it was refluxed for four hours in an al-
coholic sodium hydroxide solution. Water precipitated a product
which crystallized from ether in large clear needles, melting at 72°.
This was shown to be paraphenoxyphenylethylcyanamide,

CeH50C,;H,C,H^NCN (XV) ,
the formation being due to the opening of the ring and loss of thio-
acetic acid. This proved the position of the ethyl group at position
No. 2.

Anal. Calcd. for CigHi.ON^; N, 11.76. Found: 11.58,11.65.

For further proof the cyanamide (XV) was synthesized as fol-
lows: Some p-phenoxyphenylethylamine was made by refluxing
p-phenoxy aniline with ethyl iodide and vacuum distilling the oil,
which boiled at 213-214° at 32 m. m. pressure. This oil was treated

218 The University Science Bulletin

with cyanogen bromide and the resulting product taken up in ether.
The ether sokition was washed first with acid, then with alkali, to
remove unchanged amines and the mono substituted cyanamide.
The ether on evaporation yielded a compound which crystallized
from heptane and melted at 72°. This was shown by mixed melt-
ing point and analysis to be identical with XV.

An independent proof of the constitution of the 2-p-phenoxy-
phenyl-2-ethylamino-4-thiazolidone was carried out as follows:

2-p-Phenoxyphenyl-2-ethylamino-5-benzal-4-thiazolidone,
S — C (NC„H,OC«H,^C,H, ) NCOC = CHC.H^ (XVI)

This resulted from the condensation of the thiazolidone with
benzaldehyde in alkaline-alcoholic solution. The pure compound
melted at 176°.

Anal. Calcd. for a,H,,N,OS: N, 7.00. Found: 7.07,7.17.

Contrary to our usual experience, this was readily hydrolyzed on
boiling for thirty minutes in a strong alcohol-sodium hydroxide
solution. From the solution was isolated 5-benzal-2, 4-thiazoldione
(m.p. 240°) and an oil which with phenylisocyanate gave an urea
(m. p. 118°) which, by mixed melting point and analysis, proved
to be a-p-phenoxyphenyl-a-ethyl-^-phenyl urea (V) ; thus again
confirming the structure assigned.

2-p-Phenoxyphenylimino-3-ethyl-4-thiazolidone,

S-C(NC,H,OaH,)N(C,H,)COCH, (XIV)

As previously stated (page 5), this was obtained in three-percent
yield on the ethylation of the sodium salt of the thiazolidone (XI).
Anal. Calcd. for C,,H,,N.O,S: N, 8.93. Found: 8.88, 8.86.

It was synthesized for comparison by heating an alcoholic mixture
of a-7)-phenoxyphenyl-/3-ethylthiourea (VII), ethylchloroacetate and
pyridine. This product melted at 124° and was shown to be identical
with (XIV) , thus proving the position of the aryl and alkyl groups.

Benzyl Derivatives

2-p-Phenoxyphenyl-2-benzylamino-4-thiazolidone,

S-C (NC,H,OCeH,CH,CeH J NCOCH3 (XVII)

The sodium salt of the thiazolidone (XI) was boiled in alcohol
solution with benzyl chloride. The residue left after steam dis-
tillation was insoluble in dilute acid. After crystallization from
benzene it melted at 125-126°.

Anal. Calcd. for C^.HisN^O^S: N, 7.49. Found: 7.60,7.57.

Roberts and Dains: Isomeric Ortho 21D

That it had this constitution was proven by the fact that it
differed from its isomer 2-p-phenoxyphenylimino-3-benzyl-4-thia-
zolidone (XVIII) which melted at 108°. This was synthesized by
the action of chloroacetylchloride on a-p-phenoxyphenyl-/3-benzyl-
thiourea (VIII) in acetone-pyridine solution.

Anal. Calcd. forC^H.sNAS: N, 7.49. Found: 7.37.

It condensed in the usual manner with piperonal yielding the 3-4
methylene-dioxybenzal derivative which melted at 133°, XIX.
Anal. Calcd. for C3oH,,N„0,S: N, 5.53. Found: 5.49,5.40.

Action of Benzoyl Chloride

2-p-Phenoxyphenyl-2-benzoylamino-4-thiazolidone,

S _ C (NC.H.OC.Hg COCeHs) X — COCH., (XXII)

This, the first illustration of this type of benzoyl derivative, was
synthesized when the sodium salt of XI was treated, in water or
alcohol solution, with the calculated amount of benzoyl chloride.
An orange-colored product was obtained, which melted at 165°, when
crystallized from an alcohol-benzene solution.

Anal. Calcd. for CooH^.N.OgS: N, 7.22. Found, 7.28, 7.33.

The compound was hydrolyzed with ten percent HCl in an auto-
clave at thirty-five pounds pressure for two and one half hours, and
from the reaction mixture was isolated 2, 4-thiazoldione (m.p.
128°), paraphenoxyaniline and benzoic acid.

That the benzoyl group is at position No. 2 is indicated by its
analogy to the action of acetic anhydride on 2-phenylimino-4-
thiazolidone, which gave 2-phenyl-2-acetylamino-4-thiazolidone;
the constitution of the latter was proven by its synthesis from
a-phenyl-a-acetylthiourea and chloroacetylchloride (4).

Derivatives of the a-/8-DisuBSTiTUTED Thiourea

2-7)-Phenoxyphenylimino-3-7>-Phenoxyphenyl-4-thiazolidone,
S _ C (NCoH.OCeHs) N (CeH.OCeHj" COCH3 (XX)

This was made by boiling the diphenoxyphenylthiourea and ethyl-
chloroacetate either in alcohol-pyridine solution or in glacial acetic
acid. The crystals froln benzene melted at 131°.

Anal. Calcd. for C.^H.^N^OaS: N, 6.19. Found: 6.44,6.28.

Its 5-benzal derivative (XXI) formed fine yellow crystals from
benzene and melted at 151°. ,

Anal. Calcd. for Ca.H^.N.OaS: N, 5.61. Found: 5.40.

220 The University Science Bulletin

Thiazolidone Synthesis {Labile Para)

2-lmino-3-p-Phenoxyphenyl-4-thiazolidone,

S — C(NH) N (C.H.OCsHg) COCH, (XXIII)

! I

It has been mentioned previously that long boiling of the alcohol
solution of potassium thiocyanate and w-chloro-p-phenoxyacetani-
lide (X) gave only the stable thiazolidone XI (m. p. 184°). By
modifying the conditions, the intermediate products were readily
isolated.

The chloroacet compound (7 gms.) and potassium thiocyanate
(3 gms.) were refluxed for fifteen minutes in absolute alcohol. On
cooling the filtered solution, white crystals of the thiocyan-p-phe-
noxyacetanilide, CeH.OC.H.NHCOCHsSCN, (XXIV) separated
which melted at 106-107°.

Anal. Calcd. for CisH^oNAS: N, 9.86. Found: 9.59,9.53.

This thiocyanate could be recrystallized without change from
ninety-five percent alcohol, provided the concentration was such that
it separated out quickly. If it was of such concentration that all
stayed in solution on cooling, a rearrangement took place which
caused after fifteen hours a crystallization of long, fine, white in-
soluble needles. These were moderately soluble in hot alcohol and
melted at 132-133°. This proved to be the unstable thiazolidone
(XXIII).

Anal. Calcd. for C.sHioN^O.S: N, 9.86. Found, 9.69, 9.74.

On long standing or boiling in alcohol solution this labile form
changed to the stable form (XI). However, it was found that re-
peated crystallization of the labile form from the same alcohol solu-
tion produced a crystalline mixture melting at 122-123°. Investiga-
tion revealed these crystals to be a mixture of the stable and labile
forms. They were separated by the greater insolubility of the
stable form in hot benzene.

Reactions of the "Labile" Thiazolidone

2-Imino-3-p-Phenoxyphenyl-5-benzal-4-thiazolidone,

S — C(NH) N (CeH.OCeH^) COC = CHC.H^ (XXV)

I I

Under the influence of piperidine or sodium hydroxide in absolute
alcohol solution, the labile thiazolidone (XXII) condensed with
benzaldehyde, yielding the above benzal compound which melted
at 174°. The same product was obtained in an interesting way from
the thiocyanate (XXIV) and benzaldehyde in absolute alcohol solu-

Roberts and Dains: Isomeric Ortho 221

tion with piperidine. Here evidently tlie thiocyanate rearranged to
the labile form and then condensed to form the benzal derivative
(XXV). The melting point of this compound was lowered by re-
peated crystallization from alcohol, due to partial rearrangement to
the benzal derivative of the stable form (XII), thus giving a mix-
ture of the benzal derivatives of the stable and unstable thiazo-

lidones.

Urea Formation

a- (a-Napthvl) -^, 2- (3-paraphenoxyphenyl-4-thiazolidone) urea,
S_C(NCONHC,oH,) N (C,H,OC,hV) COCH, (XXVI)

When the labile thiazolidone (XXIII) and a-naphthyl isocyanate
were dissolved in dry benzene, the urea slowly separated. Crys-
tallized from acetone alcohol, it melted at 247°.

Anal. Calcd. for C26H,,N303S: N, 9.28. Found, 9.53.

a-Naphthylisocyanate would not react with the corresponding
stable form (XI).

a- (a-Naphthvl-/8, 2- (3-7>Phenoxyphenyl-5-benzal-4-thiazolidone)
urea,S — C(NCONHC,„H,) N (C«H,OC«H,) COC = CHC.H,

' - ! (XXVII)

This was made by refluxing the thiazolidone (XXVI) with ben-
zaldehyde in the usual manner. The light yellow needles melted at
221°.

Anal. Calcd. for C33H23N3O3S: N, 7.77. Found: 7.93.

The same compound was obtained by the action of a-naphthyl
isocyanate on the unstable benzal derivative (XXV), thus con-
firming the structure.

Benzoyl Derivatives

2-Benzoylimino-3-p-Phenoxyphenyl-4-thiazolidone,

S — C(NCOCeH,) N (aH.OCeH,) COCH, (XXVIII)

A mixture of the labile thiazolidone (XXIII), benzoyl chloride
and dilute sodium hydroxide was heated. The resulting product
crystallized from alcohol in white needles, melting at 214°.

Anal. Calcd. for C^^H.^N^OaS: N, 7.22. Found: 6.92, 7.08.

The isomeric benzoyl derivative from the stable thiazolidone had
a melting point of 165° (XXII). The constitution of this unusual
compound, the only thiazolidone thus far obtained with an acyl
group at position No. 2 and an aryl grou]) at position No. 3, was
proved as follows: a-benzoyl-/?-p-phenoxyphenyl-thiourea,

CeHgCONHCSNHC.H.OCeHg (XXIX)
was readily prepared by the action of benzoylisothiocyanate on

222 The University Science Bulletin

p-phenoxyaniline (5). When crystallized from acetone it melted at
128°.

Anal. Calcd. forC.oHigNAS: N, 8.05. Found: 8.09.

The thiourea was dissolved in acetone, and pyridine and chloro-
acetyl chloride added. After standing twenty hours the solution
was allowed to evaporate. A small yield of a compound melting at
214° was obtained from the residue. By mixed melting point this
was proven to be the benzoyl thiazolidone (XXVIII).

Derivatives of o-Phenoxyaniline

o-Phenoxyphenylthiourea. (XXX) .

This was made by evaporating a solution of the amine hydro-
chloride with ammonium thiocyanate and melted at 124° (6).

2-o-Phenoxyphenylamino-4-thiazolidone (XXXI) .

A satisfactory procedure for its preparation was to reflux, for three
hours, an alcoholic solution of the thiourea (XXX) with ethyl-
chloroacetate and pyridine. The "stable" product was light yellow
and melted at 147.5°. No unstable form was obtained.

Anal. Calcd. CisH^.N^OoS: N, 9.86. Found: 9.87, 9.95.

2-o-Phenoxyphenylamino-5-benzal-4-thiazolidone (XXXII) .
This thiazolidone was obtained from XXXI in the usual manner
without refluxing. The light-yellow crystals melted at 213°.
Anal. Calcd. for C.,.M,,'N,6S- N, 7.53. Found: 7.73, 7.93.

Heating in an autoclave at fifteen pounds for fifteen minutes with
3.5 percent HCl gave almost quantitative yields of o-phenoxyaniline
and 5-benzal-2, 4-thiazoldione, thus proving the structure of XXXI,

The yellow p-dimethylamino-benzal derivative of XXXI was
made in the same manner as the other benzal derivatives and melted
at 230° (XXXIII).

Anal. Calcd. for Co.H.^NgO.S: N, 9.88. Found: 9.97, 10.04.

The above compound XXXIII was hydrolyzed on boiling with
eighteen percent HCl, yielding o-phenoxyaniline and the orange-
yellow colored 5-jO-dimethylaminobenzal-2, 4-thiazoldione,
S — CONHCOC z= CHC,H,N (CH3) , (XXIV)

which melted at 292°.
Anal. Calcd. for Ci^Hi.N^O.S: N, 11.29. Found: 11.45.

The sodium salt of XXXI.

This could be made by dissolving the thiazolidone in sodium
hydroxide or by a rather unusual method, viz., refluxing a benzene

Roberts and Dains: Isomeric Ortho 223

solution of the thiazolidone with metallic sodium; the insoluble
sodium salt was washed with benzene and recrystallized from al-
cohol. It then melted at 64°, resolidified and again fused at 236° —
a result that would indicate alcohol of crystallization.

Analysis of product melting at 236° (XXV).

Anal. Calcd. for C.^H.^NASNa: N, 9.15. Found: 9.14, 9.21.

Analysis of product melting at 64° (XXXVI).

Anal Calcd. for C.^H.iN^O.SNa • 2C2HgOH: N, 7.03. Found:

7.08, 6.88.

2-o-Phenoxyphenyl-2-ethylamino-4-thiazolidone, XXXVII.

The above sodium salt, when dissolved in alcohol and refluxed
with ethyl iodide, yielded only one ethyl derivative, in which the
ethyl group was at position No. 2. The compound melted at 112°.

Anal. Calcd. for C^.H.eOaN.S: N, 8.95. Found: 8.85, 8.90.

The 5-dimethylamino-benzal derivative (XXXVIII) was isolated
as orange-colored crystals from benzene, melting at 210°.
Anal. Calcd. for C.^HosNgOoS: N, 9.48. Found: 9.42.

The Isomeric "Labile" Forms (Ortho)

The so-called "labile" forms of the orthophenoxyphenyl deriva-
tives were characterized by unusual stability and showed in gen-
eral the reactions of the isomeric para derivatives (XXIII).

Omega-chloro-o-phenoxyacetanilide,

CeH^OCsH.NHCOCH^Cl (XXIX)

A solution of o-phenoxyaniline in dry acetone was treated with
chloroacetylchloride and pyridine. The product, from alcohol,
melted at 79°.

Anal. Calcd. Ci.Hi.NO.Cl: N, 5.35. Found: 5.22, 5.37.

Heating this compound with potassium thiocyanate in alcohol
solution for periods varying from twenty minutes to three hours
gave only the labile form of the thiazolidone, 2-imino-3-o-phenoxy-
phenyl-4-thiazolidone, XL, and not the expected thiocyanate as in
the case of the para compound (XXIV) . This labile form crys-
tallized from alcohol or heptane as clear whitish needles and melted
at 97.5°.

Anal. Calcd. for CisHj^N^OsS: N, 9.86. Found: 9.79, 9.82.

Properties of XL. The dry thiazolidone can be heated to 130°
for twenty minutes without change, and heating for less than eight
hours produced no change. Longer than eight hours gave un-
identifiable products. Dry HCl gas gave a salt which melted at 242°,

224 The University Science Bulletin

which could be neutralized to give the original compound. The HCl
salt, when heated in glacial acetic acid with sodium acetate and
benzaldehyde, gave the benzal derivative of the stable form, melt-
ing at 213° (XXII). The benzal derivative of the labile form could
not be obtained.

Hydrolysis of XL. Heating with dilute acid usually gave o-phe-
noxyaniline, thioglycollic acid and gimimy products. However,
more or less complete conversion to the stable form XXXI resulted
in the following cases. (A) Heating at fifteen pounds pressure for
twenty minutes with HCl (2 c.c.) and water (80 c.c). (B) Dis-
solving in glacial acetic acid and heating for twenty minutes at
fifteen pounds pressure caused almost quantitative rearrangement
to the stable form (m. p. 147.5°). (C) Cooling rapidly a hot aque-
ous solution of the acid salt of the unstable form. (D) Some re-
arrangement is produced by refiuxing a short time with alcoholic
potassium hydroxide.

Thiohydantoic Acid: Hydrolysis to a thiohydantoic acid was ac-
complished by dissolving in 60 c. c. of alcohol and 10 c. c. of HCl.
After refiuxing for two hours, the solution was evaporated and from
the residue was obtained a white hydrochloride salt, melting at 129°.
It gave tests for sulfur, nitrogen and chlorine and decomposed with
alkali to phenoxyaniline and unidentified products.

Anal. Calcd. for CisH^^N^OgS ■ HCl. N, 8.27; HCl, 10.76. Found:
N, 8.29; HCl, 11.00.

These results agree with the assumption that the compound (m. p.
129°) is the hydrogen chloride salt of o-phenoxyphenylthiohydantoic
acid, C,3H,OC6H,NHC(NH)SCH,COOH-HCl (XLI).

This is in harmony with the observation of Wheeler and Johnson
(7), who obtained from the labile 2-imino-3-phenyl-4-thiazolidone, a
phenyl hydantoic acid.

Urea Derivatives

a- (a-Naphthyl) -^, 2- (3-o-Phenoxyphenyl-4-thiazolidone) Urea,
S — C(NCONHC,oH,) N (C,H,OC,H,) COCH, (XLII)

The above compound resulted from the reaction of the 2-imino
compound (XL) with a-naphthylisocyanate in benzene on standing
for several daj^s. The white crystals from an acetone-alcohol solu-
tion melted at 129°.

Anal. Calcd. for C^oH^NgOgS: N, 9.28. Found: 9.01.

When this compound was heated in alcohol (60 c.c.) and hydro-
chloric acid (10 c.c), there was isolated some gummy products,

Roberts axd Dains: Isomeric Ortho 225

di-a-naphthylurea and a fonipound containing sulphur, chlorine and
nitrogen; the analysis of which agreed with its formulation as the
hydrochloride salt of the a-naphthyl urea of o-phenoxyphcnylthio-
hydantoic acid,

CioH.NHCONC ( NHC,H,OC,H, ) SCH.COOH • HCI (XLIII)

This pure white compound melted at 172°.

Anal. Calcd. for C.eHosNsO.SCl: N, 8.28. Found: 8.41.

The naphthylurea (XLIT) was found to condense with /)-di-
methylamino-benzaldehyde, yielding a (a-naphthyl >-/:?, 2(3-o-phe-
noxyphenyl-5-/j-dimethylaminobenzal-4-thiazolidone) urea, XLIV ;
a very difficultly soluble yellow product melting at 209°.

Anal. Calcd. for C,,li,,'N,0,S: N, 9.94. Found: 9.59.

This compound on acid hydrolysis gave di-a-naphthylurea and 3-
o-phenoxyphenyl-5-7J-dimethylaminobenzal-2, 4-thiazoldione, XLV,
melting at 167°, which will be described later. (Page 226.)

The naphthyl urea (XLIT) likewise condensed with benzaldehyde
in the usual manner, yielding light-yellow crystals, melting at 227°
(LIII).

Anal. Calcd. for C33H,.,N,,X),S: X, 7.76. Found: 7.70.

This product was very stable, but on heating with eighteen per-
cent HC'l at forty pounds for two hours hydrolysis occurred. The
compounds isolated were di-a-naphthylurea and 3-o-phenoxyphenyl-
5-benzal-2. 4-thiazoldione, XLVI, which separated from benzene in
yellow crystals, melting at 178°.

Anal. Calcd. for C.^Hi^NO^S: N, 3.73. Found: 3.98.

Derivatives oe Phenylisocyanate

a-Phenyl-^, 2 (3-o-Phenoxyphenyl-4-thiazolidone I urea.
S — C (NCONHCeH;) N (CoH,OCoH„ I COCH, (XLVII)

The labile thiazolidone (XL( and phenylisocyanate gave, in ben-
zene solution, the above urea. The white crystals melted at 190°.
Anal. Calc.l. for C,,H,,NX),S: N, 10.42. Found: 10.44.

The yellow 5-benzal derivative of XLVII melted at 242°, XLVIII.
Anal. Calcd. for C,,,H,^N,.A^: N, 8.55. Found: 8.37.

When tlu' plienylurea (XLVII) was treated with alcohol (60 c.c.)
and HCI (10 c.c.) and refluxed a short time, a compound was ob-
tained which corresponded to a salt of a thiohydantoic acid. This
white compound melted at 139°, and had the probable formula,
C,H,OC„H,NHC (NHCONHCeH,) SCH.COOH • HCI (XLIX)

15—4141

226 The University Science Bulletin

Anal. Calcd. for C,,H,„N30,SC1: N, 9.81. Found: 9.90.

2-Benzoylimino-3-o-Phenoxyplienyl-4-thiazolidonc,

s'— C(NCOC,H,) N (cXoCgH;) COCH., (L)

I I

A mixture of the labile thiazolidone (XL I, benzoyl chloride and
sodium hydroxide solution gave an insoluble product which was
purified from acetone. The yellowish crystals melted at 16-1° C.

Anal. Calcd. for a^HisN AS: N, 7.22. Found: 7.36,7.44.

When o-jihenoxyaniline was treated with benzoylisothiocyanate
fine white crystals, melting at 172°, were formed, which proved to be
a-o-phenoxyphenyl-^-benzoylthiourea,

CsHgOCeH^NHCSNHCOCsH, (LI)

Anal. Calcd. for C^oHi^N.O.S: N, 8.05. Found: 8.00.

This, however, failed to give the thiazolidone (L) with chloro-
acetylchloride, and hence proof of structure as in the case of the
corresponding p-derivative was not realized.

Hydrolysis of the thiazolidone (L) with dilute acid gave the hy-
drochloride salt of o-phenoxyphenylthiohydantoic acid, which has
been previously described (XLI).

When the thiazolidone L was treated with p-dimethylaminoben-
zaldehyde in the usual manner, condensation occurred with the
formation of 2-benzoylimino-3-orthoi)henoxyphenyl-5-p-dimethyl-
aminobenzal-4-thiazolidone, LII, which melted at 233° C.

The same product was obtained by condensing the labile thiazo-
lidone (XL) and ;>dimethylaminobenzaldehyde and causing this
product to react with benzoyl chloride. The deep-orange crystals
melted at 233° C. and were proven by mixed melting points and
analysis to be identical.

Anal. Calcd. for C31H05N3O3S: N, 8.09. Found: 7.88, 7.91.

Hydrolysis in acid solution eliminated the benzoyl group and
gave 3-o-phenoxyphenyl-5-/>dimethylaminobenzal-2.4-thiazoldione,
XLV, the yellow crystals melting at 167° C.

Anal. C^lcd. for C24H.3o03N,S: N, 6.73. Found: 6.80.

SUMMARY

Substitution of 0- or p-phenoxyphenyl at position No. 2 showed
the characteristic reactions of the analogous derivative previously
studied.

The 2-imino derivatives, which are the so-called "labile" types,
were found to be more stable in the ortho series than in the i)ara

Roberts axd Dains: Isomeric Ortho 227

series. They both gave stable benzoyl and urea derivatives, which
were studied in detail, as were the corresponding substitution deriva-
tives at position No. 5.

BIBLIOGRAPHY

1. Long and Dains. C. A., 2S, 2356 (1934).

Eberly and Dains. J. A. C. S., 55, 3859 (1933) ; oS, 2544 (1936).
Davis and D.\ins. J. A. C. S., 57, 2627 (1935).

2. SuTER. J.A.C.S., 57, 2852 (1929).

3. Dains, Brewster, and Olander, Org. Synthese.?, 437.

4. Dains, et al. U. of Kansas Science Bull. 24, No. 2 (1936).

5. Douglas. J. A. C. S., 56, 719 (1934).

6. H. G. Underwood. Unpublished Thesis, JJ. of Kansas, 1931.

7. Wheeler and Johnson. Am. Chem. J., 28, 141 (1902).

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 12

Notes on the Snakes of the Genus Salvadora

HOBART M. SMITH,

Department of Zoology, University of Kansas

Abstract : A brief synopsis, with a key, is presented of the differential
characters of the six species and subspecies of the genus Sahfodora. The study
is based on thirty-three specimens, twenty-three of which are from Mexico,
representing all known forms of the genus.

THE following notes are based upon thirty-three specimens,
mostly Mexican, of six species and subspecies of Salvadora.
Alost of the material is contained in the Mexican collection of Dr.
E. H. Taylor and myself. Catalogue numbers, unless otherwise
stated, refer to specimens in this collection. The remainder of the
material studied is in the Dyche Museum of X'atural History at
Kansas University. These specimens are designated by KU.

Two rather distinct groups in the genus are apparent, one con-
taining mexicana and pulcherrima, the other containing bairdii,
grahamiae, hexalepis and virgultea. The separate identity of bairdii
is beyond question, but its relationship to grahamiae is yet uncertain.
Most recent authors have considered grahamiae and hexalepis sub-
species of each other. With reluctance I have adhered to this con-
clusion, but the matter does not seem definitely proved.

I am indebted to Dr. E. H. Taylor for assistance in the preparation
of this paper, and for making possil^le the collection of much of the
Mexican material; and to Mr. C. D. Bunker for permission to study
the material in the Dyche Natural History ^Museum.

Key to the Forms of the Genus S.alvadora

1. One preocular; tail thirty percent of total length, or more; subcaudals 121 to

139 ; three supralabials entering eye; rostral not greatly enlarged 2

Two or more preoculars; tail less than thirty percent of total length; suhcandals

82 to 103; one or two supralabials entering eye; rostral sometimes greatly en-
larged 2

2. Stripes extending the full length of body and tail pulcherrima. p. 230

Stripes broken on anterior part of body and rei)laced on neck \>y indistinct cross-

oands mexicana, p. 231

(229)

230 The University Science Bulletin

3. Anterior section of nasal separated from second supralabial ; dorsolateral dark
stripes terminating on nape, not passing over temporal region of head. .. .bairdii, p. 232
Anterior section of nasal in contact with second supralabial ; dorsolateral dark
stripes may or may not pass across temporal region to eye 4

4. Two loreals ; second supralabial separated from loreals ; nine or ten supralabials . . 5
One loreal ; second supralabial in contact with loreal ; eight, rarely nine, supra-
labials; dorsolateral dark stripes passing across temporal region to eye.

g. grahamiae, p. 234

5. Middorsal light stripe one and two half scale rows wide on fore part of body;
a single, broad, dark band on either side of middorsal light stripe, not passing
onto temporal region, but fusing generally with color of head; usually only the

sixth supralabial entering eye g. virgultea* p. 236

Middorsal light stripe three scales wide on fore part of body (wider on neck) ;
two dark bands lateral to middorsal light stripe, narrowly separated from each
other; a distinct, dark, temporal stripe, continuous with upper dark band on
body; usually two or more supraoculars enter eye g. Itexalepis, p. 235

Salvadora pulchcrrima (Cope)

(Plate XXII)

One specimen, a male (No. 4669), was collected near San Ricardo,
Chiapas, September 2, 1936. It was found moving about the base of
a red sandstone cliff.

The essential features of scutellation are: parietals truncate pos-
teriorly; frontal emarginate on sides, wider anteriorly than poste-
riorly, slightly wedged between the parietals posteriorly, its length
on a median line somewhat greater than that of the median suture of
the parietals; prefrontals wider than long, extended onto loreal re-
gion; internasals as long as broad, rounded anteriorly, the median
suture between them about two thirds the length of the median
suture between prefrontals; maximum length of prefrontals slightly
greater than that of internasals; rostral nearly one and one half
times as long as broad, wedged posteriorly between internasals; an-
terior section of nasal subequal in size to posterior section, not in
contact with second supralabial ; dorsal border of naris formed by
internasal; loreal single, narrow, elongate; one large prcocular, ex-
tending dorsad and nearly completely separating jirefrontal from
supraocular; preocular in contact only with third supralabial; supra-
ocular wider posteriorly than anteriorly, its greatest transverse diam-
eter somewhat less than that of frontal; two postoculars, the lower
somewhat smaller than upper; sujiralabials 8-8, the second and
third in contact with loreal, the third, fourth and fifth entering eye
(third very narrowly); temporals 2-2 on each side; infralabials
11-11, the first pair in contact on the median ventral line; two pairs
of genials, the length of the first pair, on the median ventral line,
slightly less than that of second pair; scales of second pair of genials

* Characters as proposed by Bogert (Bull. Southern Calif. Acad. Sci., vol. 34, part 1,
1935, pp. 88-94). with the exception of the absence of a temporal stripe.

(

Smith: Genus Salvadora 231

separated by two small median scales anteriorly, by three scales
posteriorly.

Dorsal scales smooth, with two apical pits, in 17-17-13 rows;
gastrosteges 199; iirosteges 138. The total length is 1,056 mm.;
tail, 345 mm. ; the latter is 32.7 percent of the total length.

The color pattern is quite distinctive. Most of the head scales are
edgetl with black; general ground color of head light olive; ground
color of body light olive anteriorly, fading to cream posteriorly;
four distinct, black stripes on neck; the two medial stripes begin
three scales back of the parietals; they are two scales wide on the
nape and are separated medially by a light stripe five scales wide;
a very short distance posteriorly they are separated by a wddth of
three scales, diminishing to a width of one scale one fourth the dis-
tance from the snout; on the tail the median light stripe becomes
narrower, very light brown, and occupies parts of two adjacent scale
rows; the black stripes on either side of the middorsal light stripe
become dark brown near the middle of the body and progressively
lighter posteriorly; near the middle of the body, the scales near the
edges of the dorsolateral dark stripes are dark-edged, while pos-
teriorly only the upper edges of the scales on the lateral border of
the dark stripes are black.

The lateral black stripes extend from the posterior border of the
eye through the temporal region to the neck, where they are broken
by a light area about the length of four scales; they continue
posteriorly from the neck, where they are about two half scale rows
wide, to the tail, becoming brownish on the posterior fourth of the
body; over most of the length of the body the lateral stripes are
one and one half scale rows wide, including the upper half of the
lateral scale rows.

The belly and the ventral half of the first dorsal scale row are
immaculate white; the chin and all except the upper edges of the
supralabials are white.

Salvadora mexicana (Dumeril and Bibron)

Thirteen specimens are available, from the following localities:
Colima: Manzanillo (No. 4676). Michoacdn: Hacienda El Sabino
(Nos. 4680, 5265-8). G-uerrero: Mexcala (Nos. 4673, 5269, 15424) ;
1 mi. N. of Organos, S. of El Treinte (No. 4674) ; El Treinte (No.
5270) ; Palo Blanco, S. of Chilpancingo (No. 4675).

This species agrees with pulcherrima in most respects, but has a
somewhat larger rostral and the supralabials are nine on each side

232

The University Science Bulletin

instead of eiglit. The anterior section of the nasal is separated from
the second supralabial in all specimens. The fourth, fifth and sixth
.supralabials border the orbit in all except one side of one specimen,
in which the fourth is narrowly excluded by contact of the pre-
ocular and fifth supralabial. The dorsal scale formula is 17-17-13
in all. The infralabials are eleven except on one side of one speci-
men, and on both sides of another, in whicli there are ten.

Measurements (in mm.) and .scale counts of S. uuxicrma

Number.

Sex.

Ventrals.

Caiidals.

Total
length.

Tail.

Percent
tail of
total.

5266

9
9
d'
cf
d'
&
&

&

192
186
189
181
186
188
191
187
185
185
187
188
186

129
137
121
132
141
137
131
123
131
128
139
134
132

555

918

656

760

797

841

1,032

1,049

1,063

1,092

1,140

1,274

1 ,330

174
309
210
258
274
289
341
337
341
362
386
425
432

31.4

5269

33.7

4676 ...

32.0

4673

33.9

4675

34.3

15424

34.4

5267

33.0

5268

32.1

4674

32.1

5265

33.2

5271

33.9

5270

33.4

4680

32.5

The species differs widely from others of the genus in coloration.
The striped pattern is replaced on the anterior third or fourth of
the body by a speckled pattern; on the neck the dark spots are
arranged in irregular, indefinite cross-bars about six in number.
Each dorsal head scute has a large, light area in the middle, sur-
rounded by a darker area around the edge of the scale. A dark
spot occurs at the lateral anterior edge of each caudal scute.

Salvadora bairdii (Jan)

Seven specimens are available, from the following localities: Vera
Cruz: It/o mi. W. of Acultzingo (Nos. 4668-9). Puebla: 12 mi. N.
of Tehuacan (Xo. 5264). Guanajuato: San Felipe (No. 4670).
Jalisco: Magdalena (Nos. 4670A, 467] l. Michoacdn: Truapan (No.
4672).

The species is similar to grahamiac grahamiac, witli which it has
been synonymized by some authors, in most characters of cephalic

Smith: Genus Salvadora 233

sciitcllation. There are eight supralabials; usually nine iiifralabials
(10 on one side in two specimens) : two preoculars; two postoculars;
lower preocular wedged between third and fourth supralabials; sec-
ond and third supralabials bordering loreal; latter single, deep;
fourth and fifth supralabials enter eye. However, there are three
definable differences in the cephalic scutellation of grahamiae gni-
hamiae and bairdii. In the latter the anterior part of the nasal is
not pushed back by the rostral, and is separated from the second
supralabial by contact of the first supralabial and the posterior part
of the nasal. In g. grahamiae the anterior section of the nasal is in
contact with the second supralabial, presumably due to the enlarge-
ment of the rostral. In g. hexalepis the rostral is still further en-
larged, resulting in a broader contact of the two scales mentioned.

Secondly, the rostral is not enlarged in bairdii as in g. grahamiae,
and the lateral edges are but very slightly free. The outline of the
anterior face of the rostral, in dorsal or ventral profile, is rounded
in bairdii, nearly straight in g. grahamiae.

Thirdly, the second pair of genials in bairdii are separated
throughout their length, as is usually the case in g. hexalepis; in g.
grahamiae the scales usually are in contact throughout most of their
length, being separated only posteriorly.

Further, the pre- and postoculars seem to vary less in bairdii than
in g. grahamiae. There are invariably two pre- and two postoculars
in the specimens examined of bairdii, while in six specimens of g.
grahamiae the preoculars are 2-2, 2-2, 2-3, 2-3, 2-3. 3-4. and there
are three postoculars on one side in one.

In ventral and caudal counts and in tail-total length proportions
no differences are discernible.

The color pattern of bairdii is essentially similar to that of g.
grahamiae, but there are two definite and constant differences. The
dorsolateral dark stripes in bairdii are distinctly black-edged medi-
ally except on the posterior part of the body, not in g. grahamiae.
In the latter the dorsolateral dark stripes pass distinctly onto the
head, through the temporal region to the eye, while in bairdii the
stripes terminate on the nape just posterior to the parietal and
temporal regions.

Further, the narrow, lateral dark stripe in bairdii extends farther
forward than in g. grahamiae. The median dorsal light stripe in
the latter species is one and two half scale rows wide on the body
near the anus, while in all but one specimen of bairdii the dorsal
stripe has nari'owed to the width of one scale row at this point.

234

The University Science Bulletin

In bairdii dark blue, irregular, vertical lines passing from the
lateral dark line to the ventrals, frequently are visible on the neck.
These are not or scarcely discernible in g. grahamiae.

Salvadora grahamiae grahamiae and bairdii are undoubtedly
closely related — apparently more closely than g. grahamiae and g.
hexalepis, as indicated by the greater number of definable differ-
ences between the latter two than between the former two. Upon
the basis of present knowledge, however, it can merely be stated
that bairdii and g. grahamiae are distinct from each other. The
degree of difference between related forms is no criterion for de-
termining whether the forms are species or subspecies. Two sub-
species of a single species may have more obvious, more numerous
and more easily defined differences (e. g., Sceloporus m. magister and
m. rufidorsiim) than two closely related forms known to be distinct,
specific entities through the existence of other evidence (e. g., Scelo-
porus m. magister and c. clarkii). In the absence of evidence in the
form of geographic trends of variation indicative of a blending of
differential characters in an area between the known ranges of the
two forms in question, I prefer to retain them as separate species.

Measurements (in mm.) and i^cale counts of S. bairdii

Number.

Sex.

Ventrals.

Caudals.

Total
length.

Tail.

Percent
tail of

total.

4672

9
9

194
191

90
94

261
297

54
70

20.7

4668

23.6

5264

9

206

95

353

81

22.9

4670

9

209

98

663

163

24.6

4670A .

9

197
185

688
254

4669

100

61

24.0

4671

&

186

95

694

182

26.2

Salvadora grahamiae grahamiae Baird and Girard

Six specimens are in the collections, from the following localities:
Nuevo Leon: Spring three miles west of Sabinas Hidalgo (No.
4677). Texas: Arroyo El Salado, 13 m. SE of Rio Grande City,
Starr Co. (No. 4678) ; Lytle, Atascosa Co. (KU 116'68). New Mex-
ico: Carlsbad Caverns, Eddy Co. (KU 8380) ; Santa Rita, Grant
Co. (KU 2142). Arizona: Montezuma Canyon, Huachuca Mts.
(KU5467).

Supralabials eight, except in one in which nine are present on one
side; infralabials nine, except in one in which ten occur (both sides) ;

Smith: Genus Salvadora

235

preoculars 2-2, 2-2, 2-3, 2-3, 2-3, 3-4; postoculars two, except on one
side of one, where three occur; lower preocular inserted between
third and fourth supraLabials, except on one side of one (between
fourth and fifth) ; second and third supralabials in contact with
loreal; fourth and fifth supralabials usually entering eye (fifth and
sixth on one side in one) .

The anterior section of the nasal is in contact with the second
supralabial in all. In two specimens, the second pair of genials are
separated throughout their length.

Measurements (in mm.) and scale counts of S. g. gtuhamiae

Number.

Sex.

Ventrals.

Caudals.

Total
length.

Tail.

Percent
tail of
total. '

11668

9
&

189
185

92

94

92

103

97
85

918
516
724
752

847
909

222
135
188
202
218
222

24.2

4677

2142

26.2
26.0

8380

5467

4678

191

184
187

26.9
25.7
24.4

Salvadora grahamiae hexalepis (Cope)

Five specimens, from the following localities: Sonora: 50 kilom.
S. ofNogales (No. 15426). Arizo7m: Roosevelt Dam, Gila Co. (KU
8429) ; Mohawk, Yuma Co. (KU 6997) ; Phoenix, Maricopa Co. (KU
8488) ; Tucson, Pima Co. (No. 4679).

Dorsal scales in 17-17-13 rows, as in g. grahamiae and bairdii;
supralabials 9-9 in two, others 9-10, 10-10; infralabials 9-10, 10-10,
10-11, 11-11; preoculars two in all; postoculars 2-3 in one, 2-2 in
others; anterior section of nasal in contact with second supralabial
in all ; loreals 2-2 in two specimens, 2-3 in one, 3-3 in two ; third and
fourth supralabials in contact with loreals, except on one side in two
specimens, in which the third, fourth and fifth supralabials are in
contact with the loreals ; lower preocular wedged between the fourth
and fifth supralabials except on one side in two specimens, in which
it is wedged between the fifth and sixth. The supralabials entering
the eye are as follow^sc 5-6-7. 7; 6,6; 6,6; 0,5. In the latter speci-
men, the sixth supralabial, which should enter the eye, is split trans-
versely. A subocular, split usually from the upper edge of the fifth
supralabial, is present on both sides in three, on one side in one
(head crushed in other specimen). The second ]iair of genials are in
contact in one specimen.

236

The University Science Bulletin

The subspecies differs from g. grahamiae in numerous characters:
number of supralabials (nine or ten in hexalepis, eight or rarely nine
in grahamiae) ; number of infralabials (nine to eleven, usually
eleven, in hexalepis; nine or ten, usually nine, in grahamiae) ; num-
ber of loreals (two or three in hexalepis, one in grahamiae) ; second
supralabial (in contact with loreal in grahamiae, not in hexalepis);
rostral wider in hexalepis; anterior section of nasal more broadly in
contact with second supralabial ; second pair of chinshields more fre-
quently separated in hexalepis; and ventral scales more numerous
in hexalepis.

Measurements (in nun.) and scale counts of <S'. g. hexalepis

Number.

Sex.

Ventials.

Caudals.

Total
length.

Tail.

Percent
tail of
total.

8488

6997

4679

9
9
&

201
207
201

82
9.5
86

709
826
684

154
182
163

21.7
22.0
23.8

Salvadwa grahamiae virgultea Bogert

One specimen examined, from Escondido, San Diego county, Cali-
fornia ( KU 8487 1 . It is a young female, agreeing in most characters
with Bogert's description (Bull. Southern Calif. Acad. Sci., Vol. 34,
1935, part 1, pp. 88-94). The dorsals are in 17-17-13 rows; ventrals
201; caudals 84; supralabials 9-9; infralabials 11-11; preoculars 2-2;
postoculars 2-2; loreals 2-2; third and fourth supralabials touch
loreals; lowTr preocular between fourth and fifth supralabials; fifth
and sixth supralabials enter eye on one side, sixth on other; sub-
ocular present on one side; total length 277 mm.; tail 59 mm.; ratio,
tail to total length, .213.

In addition to the characters pointed out by Bogert, it appears
that g. virgultea differs from g. hexalepis also in lacking a distinct,
dark temporal stripe. It agrees in this respect with hairdii.

Smith : Genus Salvadora

237

platp: XXII

Salvadora pulchcrrima. San Ricardo. Chiapas. Total length, 1056 mm.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXVJ June 1, 1938 [No. 13

Miscellaneous Notes on Mexican Snakes

EDWARD H. TAYLOR and HOBART M. SMITH,

Departiiient of Zoology, University of Kansas

Abstract : The following spocie.? of Mexican snakes are discussed : Loxoce-
mus bicolor Cope; Diadophis regalis dougesii (Villada) ; Conopsis jrontalis
(Cope); Xenodon angustirostris Peters; Geophis semidoUatus Dumeril and
Bibron; Geophis blanchardi sp. nov. (related to dubiics and chnlybaeus) ; Enu-
lius unicolor (Fischer); Enidius s-iimichra-sii Bocourt ; Storcria dekayi (Hol-
brook) ; Storcria storerioidcs (Cope); Ninia diademata Baird and Girard;
Chersodromus liebmanni Reinhardt; Conophis concolor Cope; Conophis vit-
tatus Peters; Tantilla rubra Cope; Tantillo bocourti (Giinther) ; Tantilla cal-
amarina Cope; Stenorhina dcgcnhardti apiata Cope; Stcnorhina degenhardti
quinquelineata (Hallowell), Clelia clelia (Daudin).

THE following notes are based on specimens collected, for the
most part, by tlie authoi's, in Mexico.

Lo.vocemus bicolor Cope

Loxocemus bicolor Cope, Proc. Acad. Nat. Sci. Philadelphia, 1861, p. 70 (type description;
type locality. La Union, Salvador).

A single specimen of this species (EHT-HMS No. 4574, 2 mi.
north Xaltianguis, Guerrero, Mexico) was collected on the highway
at night during a downpour of rain ; lights from my car disclosed it
crawling across the road.

Rostral large, upturned, part visible above three fourths of the
distance from frontal; internasals about half area of prefrontals,
narrowed medially; prefrontals bordered by three labials; no loreal,
frontal longer than its distance from end of the snout ; parietals small,
separated posteriorly by an occipital shield; one large preocular
broadly in contact with the frontal; a supraocular, small, displaced
backwards by the upper extension of the preocular; three post-
oculars; temporals 3+6 + 6+7; upper labials 11,11, the seventh and

(239)

240

The University Science Bulletin

eighth on the left side, the sixth, seventh and eighth on the right side
are very small, as if the normal scales had segmented since the
temporals above them are enlarged; fourth and fifth labial enter
orbit; 13, 14. lower labials, three touching the anterior chinshields,
which are more than double the size of the second ])air; a large scale
borders lower labials 4 to 7, inclusive, on left side, broken in two
parts on right side; 13 scales between chinshields and first widened
ventral; ventrals, 256; subcaudals 44; two pairs of divided preanals;
scale formula 44. 36, 34, 34, 28, 26. Dark lavender brown, the head
darker; the color likewise darker on the underside of head; ventral
surface with considerable brownish pigment, the back part of each
ventral light. An anal cream spot; a cream-colored area on snout
below nostril.

Diadophis regalis dougesii (Villada)

Diadophis piincfatiis Dougenii Mllada, La Naturaleza. Ill, 1874-76 (1876), pp. 226-230. pi.
(lower figuii' in color, and A-C). Type description; type locality "potreros de Balbuena"
(east of Mexico City).

A single specimen in the collection (EHT-H]\IS No. 4587 5 ; near
Corones, Guanajuato, Aug. 12, 1932. Taylor and Smith). It was
taken crawling across the road early in the afternoon.

Ventrals, 203; anal divided; caudals, 49; upper labials, 7-7;
lower labials, 8-8; preoculars, 2-2; postoculars, 2-2; temporals.
1+1+1, 1+2+2; labials touch chinshields, 5-5; labials enter eye,
3, 4; scale formula, 21, 17, 17, 15, 15. Measurements (in mm.):
Total length, 504; tail, 87; body width, 11; head width, 10; head
length, 14.6; jiarietal length, 6; parietal to tip of snout, 5.6; frontal

Fig. I. Diddophi.'i n(jalis doiu/c.vl (\'illa(ln) EHT-HMS No. 4587; Corones,

Guanajuato, Mexico. X3.

Taylor and Smith: Mexican Snakes 241

length, 3.8; frontal width, 2.5; diameter of eye, 2.05; eye to nostril,
2.5. Maxillary teeth, 9-10, followed after an interspace by a large
solid tooth. Scale preceding anal plate divided.

This form is a much heavier snake than the Arizona form, the
head and body are distinctly wider, the head proportionally shorter.

The color is slaty gray to brownish gray, darker posteriorly, each
scale with numerous silver flecks when seen under a lens. Head
same, but darker; orange neck band about one and one half scale
rows wide; outer edge of each ventral blackish, the color extending a
third of the way across posterior part of ventrals. Outer row of
scales without orange save the first four or five scales; labials with
an orange line meeting on snout; many labials with spots or dark
edges; chin and ventral surface orange with numerous small black
spots, usually two on each ventral besides larger ones on outer edge
of scale; under tail red-orange.

This specimen has nearly the same ventral count as that given for
the type (202) ; the number of caudals is larger since the tail of the
type specimen is incomplete. The type measured 650 mm. in length ;
the width of the body is 15 mm.

Conopsis frontalis (Cope).

(Plate XXIII, fig. 3)

Tallica frontalis Cope, Proc. Acad. Nat. Sci. Philadelphia, 1864, p. 167 (type description;
type locality, Coliiiia).

Ficimia otivacea (part.) Peti^rs, Mon. Akad. Wiss. Berlin, 1869, p. 875.

Geagras frontalis Cope, Journ. Acad. Nat. Sci. Philadelphia, (2), VIII, 1876, p. 142
(States: "Toluca frontalis Cope, from Colima, is congeneric with this specius [Geagras redim-
itus] in all technical characters."); Amer. Nat., 1884, p. 163; Bull. U. S. Nat. Mus., No. 32,
1887, p. 82 (Colima and Guadalajara).

Ficimia frontalis Ciarman, Mem. Mus. Comp. Zool. Harvard, VIII, No. 3, 1883, p. 82.

Pseudoficimia pulchra Bocourt, Mission Scientifique au Mexique et dans I'Amerique Centrale,
Kept. Livr. 9, 1883, pp. 572-573, pi. XXXV, fig. 12, and 12a-12e (type description; type
locality, "Mexique").

Pseudoficimia frontalis Gunther, Biologia Centrali-Aniericana, Reptiles and Batrachia,
May, 1893, p. 96 (Ventanas, Durango, and Presidio, Sinaloa) ; Cope, Ann. Rept. U. S. Nat.
Mus., 1898 (1900), pp. 945-946.

Contia frontalis Boulenger, Cat. Snakes British Mus., 2d Ed. 1894, p. 270; Werner, Zool.
Jahrb., 57, 1929, pp. 147, 149.

Conopsis frontalis Amaral, Mem. Inst. Butantan, IV, 1929, p. 182.

From the above synonymy it is obvious that opinions as to the
generic relationship of this species have differed. The most reason-
able association is with Conopsis, since its most closely related
species is Conopsis nasus.

Two specimens of this form (EHT-HMS, 5203, 5204) were col-
lected at Hacienda El Sabino, about 25 Ion. south of Uruapan,
]\Iichoacan, by Don Julio Raymond Bresson, who presented the
specimens to Smith.
16—4141

242 The University Science Bulletin

The following data are from Nos. 5203 $ and 5204 j , respec-
tively. Scale fonnula, 22, 17, 15, 17; 28, 17, 17, 17; ventrals, 163,
158; subcaiidals, 34, 39; upper labials, 7-7, 7-7; lower labials, 7-7,
7-7; temporals, both 1+2+3; preoculars, 1-1, 1-1; postoculars, 1-1,
1-1; anal divided; labials touch chinshields, 3-3; 3-3; total length,
517, 450 mm.; tail, 72, 73 mm.; frontal, length, width, 6'.4X4.3,
5.6X3.8 mm.; parietals, length, width, 6.5X5; 5.9X4 mm.; frontal
to end of snout, 6, 5 mm.; rostral visible above, 2.4, 2 mm.; eye
diameter, 2.8, 2.8; eye to nostril, 3.9, 3.7. Spots on body, 40, 36; on
tail, 12, 13.

The nasal is divided below the nostril only; fourteen maxillary
teeth, the outer faces showing slight groovelike depressions. Ground
color grayish-brown with a narrow, median, cream line interrupted
by the darker brown, blackish-edged spots, the outer edges of which
extend as narrow vertical lines to the third scale row; two rows of
small, black flecks on first and second scale rows; below, cream;
yellow on throat. A few subcaudals blackish.

Xenodon angustirostris Peters

(Plate XXIII, fig. 4)

Xmodon angu>stirostris Peters, Monatsb. Akad. Wiss. Berlin, 1864, p. 390 (type descrip-
tion; type locality, Veragua, Guatemala); idem. 1873, p. 607 (three specimens "Camaron,
Provinz Chirique") ; Cope, Journ. Acad. Nat. Sci. Philadelphia, (2), VIII, 1876, p. 141
(Sipurio, Costa Rica), p. 157 (Camp Mary Caretta, Nicaragua); and Bull. U. S. Nat. Mas.,
No. 32, 1887, p. 76; Glinther, Biologia Centrali-Americana, Rept. Batr., Feb. 1894, p. 114.

Xenodon sp. MuUer, Verh. Nat. Ges. Basel VI, 1878, pp. 663-666 "Costa grande gegend
von Mazatenago, Guatemala."

Xenodon rhabdocephalus (part.) Giinther, Biologia Centrali-Americana, Rept. Batr., Feb.
1894, p. 114, Amula, Guerrero; British Honduras; San Geronimo, Guatemala; Gadow, Proc.
Zool. Soc. London, June 6, 1905, p. 233.

Xenodon cohibrinus Boulenger, Cat. Snakes British Mus., 2d Ed., II, 1894, pp. 146, 147,
(part.); Stuart, Mus. Zool. U. of Michigan, Misc. Publ., No. 29, Oct. 1, 1935 (3 spec. La
Libertad, Guatemala).

Ophis = (Xenodoii) cohibrinus Wettstein, Sitz. Akad. Wiss. Wien, abt. 1, 143 Bd. V2 Heft,
1934, p. 35 (Costa Rica).

We are uncertain as to what name is the proper one to apply to
the species of Xenodon occurring in Mexico. Giinther loc. cit. (Feb.
1894) placed specimens from Guatemala, British Honduras and
Mexico under the name Xenodon rhabdocephalus Wied, and recog-
nized Xenodon angustirostris Peters as a distinct species without
having seen a specimen. He believed that Xenodon bertholdi fig-
ured by Jan {loc. cit.) is not from Mexico, but from Brazil, and
identical with Xenodon cohibrinus Giinther.

Boulenger, 1894, referred Xenodon rhabdocephalus {part.) Giinther,
bertholdi Jan, angustirostris Peters and bipraeocularis Cope to the
synonymy of Xenodon cohibrinus Giinther, referring to it sixteen

Taylor and Smith: Mexican Snakes 243

specimens from localities ranging from Para, Brazil, to Amula,
Guerrero, Mexico.

Cope (1876, loc. cit.) referred specimens from Central America
to Xenodon angustirostris Peters (as did Bocourt, loc. cit.). He
regarded this form as a subspecies of Xenodon severus Linne.

Schmidt (Smithsonian Misc. Coll., 89 No. 1, 1933, p. 16) re-
ferred Panama specimens, which he examined, to Xenodon colu-
brinus. These specimens, four females have combined ventral —
subcaudal counts ranging from 191 to 198; Boulenger's specimens
from Panama have totals of 193; his specimens from Columbia
range from 190 to 202. Stuart {loc. cit.) referred three specimens
from La Libertad, Peten, Guatemala, to the same species. Those
specimens have total counts of 167 and 171.

Granted that specimens from Panama and Columbia are cor-
rectly referred to coluhrinus, we find that the total counts range
from 190 to 202, with an average of 194. Those from localities to
the north of Panama vary between 167 and 188, the average being
175. The type of Xenodon angustirostris has a total count of 188
and is approached by a specimen from Guatemala with 183. X.
bertholdi is probably a distinct species.

Of the specimens listed by Boulenger and Schmidt, all are young
or females. Stuart lists two males, one of which has the lowest
ventral count (124) and at the same time the lowest ventral-sub-
caudal count (167); the ventral count for the second male is 125
(subcaudal count not given).

A single specimen in our collection was collected by Dyfrig McH.
Forbes at Potrero Viejo, near Cordova, Veracruz, Mexico, and is,
we believe, the most northern record. This specimen (No. 5207 $ )
yields the following data: Scale formula, 23-19-19-17; ventrals, 123;
anal, single; subcaudals, 35; preoculars, 1-2; postoculars, 2-2; tem-
porals, 1+2+4; four labials touch the first chinshields, which are
much larger than the posterior; 15 broad bands on body, 3 on tail;
frontal length, 5.8 mm.; width, 5.5 mm.; parietal length, 6.9 mm.;
width, 5.7 mm.; frontal to tip of snout, 8 mm.; prefrontal length, 4
mm.; width, 4.2 mm.; portion of rostral visible above one third of
distance from frontal to tip of snout.

The color is grayish since the scales have been shed, the edges of
tlie spots nearly black, while the areas between the spots are clove
gray; chin yellowish; underside of tail unspotted. Belly dull cream,
flecked with very numerous ashy-brown spots; on outer edge of ven-

244 The University Science Bulletin

trals a very indistinct row of cream spots separated by darker color-
ation. The figure (PL 23, fig. 4) shows arrangement of the markings.
We are, at least temporarily, considering this Mexican Xenodon
under the name angustirost't'is Peters, despite the fact that it has a
considerably smaller count of ventrals and subcaudals.

Geophis semidoliatus Dumeril and Bibron

Eighteen specimens, three from Cordova, Veracruz (Nos. 5154-6,
collected by H. R. Roberts) and fifteen from Tlilapam, Veracruz
(Nos. 5157-71, collected by E. H. Taylor).

The specimens present the following variation in scutellation:
dorsal scales smooth in both sexes, in 15-15-15 rows; ventrals, 136
to 157 in males, 144 to 169 in females; subcaudals, 22 to 25 in
females, 23 to 28 in males. Five supralabials (four on both sides
of one, in which the fourth and fifth are fused); six infralabials;
third supralabial enters eye; no preocular; postoculars, 1-1; supra-
ocular present, small; two pairs of genials, the anterior much the
larger; anal entire.

The dark bands on the body (excluding head) vary between 17
and 32 in number; four to six are present on the tail. In most
specimens two or more of the bands are fused on the middorsal
line ; in some as many as six consecutive bands are fused. The dark
bars are usually three or four scales long, and extend laterally a
varying distance — in some specimens, terminating on the fourth
scale row, but usually on the first or second; none of the bands
extend onto the ventral surface; they are separated on the mid-
dorsal line usually by a width of two scales, laterally by a width
of three or four. The dark bands are ultramarine in young speci-
mens, and the interspaces are pink {fide E. H. Taylor's field notes),
while in larger specimens the bands are very dark blue-black, the
interspaces orange-red.

The light band across the temporal and parietal regions varies
in width from one fifth to three fourths the length of a parietal;
the broader bands have concave anterior edges.

The specimens agree with Boulenger's description (Cat. Snakes
Brit. Mus., II, 1894) except that the minimum in ventral and caudal
scale counts is lower and the maximum ventral count is less than
indicated by that author. The first maxillary tooth is posterior
to the naris, about on a level with posterior edge of the nasal.

Taylor and Smith: Mexican Snakes

245

Measurements (in mm.) and scale counts of Geophis semidoliatus

Number.

Sex.

Ventrals.

Caudals.

Total
length.

Tail
length.

5171. .

&

136
139
141
139

24
25
28
23

126
138
1.57
161

14.0

5165

14.3

5164

17.2

5160

16.0

5162

148
146

25

28

167.5
251

17.5

5169

30.5

5154

&

157

26

257

26.0

5166

&

145

25

269

28.0

5163

&

144

22

123

12.0

5167

&

154

22

129

12.1

5157

9
9

159
161

23
22

131
1.58

12.0

5170

14.1

5156

9

168

24

160.5

13.2

9
9
9
9
9

169
157
1.52
152
1.59

22
24
25
23
23

169
266
287
295
310

14.0

5158

25.0

29.5

5159

27.0

25.6

Geophis blanchardi sp. nov.

Holotype. EHT-HMS. No. 5194; collected about two miles
southwest of Acultzingo, Veracruz, Aug. 14, 1936, by E. H. Taylor.

Paratypes. EHT-HMS, Nos. 5479-5483; topotypes.

Diagnosis. A small snake, grayish-blue above, black and orange
below. Internasals and prefrontals distinct; nasal divided; an
elongate loreal; no preocular; small supraocular; one postocular;
fifth labial broadly in contact with the parietal; one large temporal;
ventrals, 155-162, and undivided; 30-40 subcaudals.

Description of type. Rostral small, not reaching the dorsal sur-
face of the head, distinctly broader than high; internasals small,
broader than long, one third to one fourth the size of the pre-
frontals; the combined^ width equal about one half of the width
of the prefrontals in front of eye, the suture between them equal to
half of that between the prefrontals; prefrontals very large, entering
the eye; frontal four-sided, the anterior angle very obtuse, the sides
converge to form a posterior angle little less than a right angle;
frontal longer than its distance from the tip of the snout; supra-

246

The University Science Bulletin

A B

Fig. 2. Geophis blanchardi sp. nov. EHT-HMS, No. 5194; type, Acultzingo,
Veracruz. Actual width of head, 6 mm.

oculars small, triangular, scarcely larger than the postocular;
parietals elongate, longer than their distance from the tip of the
snout; nasal divided into two pieces of nearly equal area, the nostril
in the anterior; loreal much elongated, its length nearly twice its
width; eye small, little more than half the length of loreal; six
upper labials in the following ascending order of size: 1, 2, 4, 3,
6, 5, the fifth far the largest, broadly in contact with the parietal;
the third and fourth enter orbit ; a single elongate temporal behind
the fifth labial; mental broader than long, not as wide as rostral;
first pair of chinshields much larger than posterior; first four ven-
trals small, equal-sized, the first touching the second pair of chin-
shields, seven lower labials in the following order of size: 2, 7, 6,
1, 5, 3, 4; three lower labials touch first chinshield; scale formula,
17-17-17; ventrals, 162, anal single; subcaudals (paired), 130; total
length, 388 mm.; tail, 46 mm.; head width, 6 mm.; head length,
10 mm.; head to end of parietal, 9 mm.

Color. Above gray-blue to blackish (where scales are shed).
Below deep orange and black, the black color forming angular spots
with an average length of three scales; these reach the middle of
belly and usually alternate with an orange spot of about equal

Taylor and Smith: Mexican Snakes 247

size; occasionally the black spots form a band across belly; chin and
neck (below) orange; lip ultramarine; a whitish spot on the edge
of fourth labial.

Remarks. The relationship is with Geophis chalybaea and Geo-
vhis dubius. From the former it differs in the entirely different
ventral coloration and in having a larger series of ventral scales;
from G. dubius it differs in having a single postocular, a different
lateral and ventral coloration, and a smaller series of subcaudals.

The species is named for the late Dr. Frank Blanchard in memory
of his excellent contributions to American herpetology.

Enulius unicolor (Fischer)

Geophis unicolor Fisher, Abh. d. natunviss. Vereines zu Bremen, vol. 7, 1881, p. 227
(type locality, "Mexico").

Leptocalanius uiiirnlor Cope, Proc. Amer. Philos. Soc, vol. 22, 1885, p. 178; Gunther,
Biol. Cent. Amer., Kept, and Amph., 1893, p. 100.

Two specimens, both females, are available, one (No. 5237) col-
lected four miles south of Cuernavaca, Morelos, the other (No.
5238) collected twelve miles south of Puente de Ixtla, in Guerrero.

The dorsal scales are in 17 rows; ventrals and caudals, respec-
tively, 192 and 93 in No. 5238, 200 and 102 in No. 5237. Third and
fourth supralabials enter eye; seven supra- and seven infralabials;
no preocular; two postoculars; temporals, 1-2-3; three scales be-
tween the single pair of enlarged chinshields and first enlarged ven-
tral; anal divided; scale preceding anal also divided; total length
and tail length, respectively, 300 mm. and 93 mm. in No. 5238, 164
mm. and 46 mm. in 5237.

Dorsal color, light tan in the larger specimen, very slightly darker
medially, the color reaching to the second scale row. In the smaller
specimen the ground color is dark olive. Belly white in both speci-
mens.

Both specimens were found under stones.

Dr. E. R. Dunn {in litt.) has called our attention to the fact that
Enulius* has priority over Leptocalamus.'\

Enulius suniichrasti Bocourt

Enulius sumichrasti Bocourt, Miss. Sci. Mex., Kept., Livr. 9, 1883, p. 538, pi. 31, fig. 6
(type locality, "Isthmus of Tehuantepec") ; Boulenger, Cat. Snakes Brit. Mus., II, 1894, p.
250; Werner, Zool. Jahrb., vol. 57, 1929, p. 143.

Enulius m.iiriniis Bocourt, Miss. Sci. Mex., Rept., Livr. 9, 1883, p. 537, pi. 35, fig. 6
(Tehuantepec specimens).

Geagras longicaudatus Cope, Amer. Nat., 1884, p. 162 (new name for Enulius murinus
Bocourt).

* Cope, Proc. Amer. Philos. Soc, vol. 11, 1871, p. 558 (type jnurinus, from Chinandega,
Nicaragua).

t GUnther, Ann. Mae. Nat. Hist.., Ser. 4, vol. 9, 1872, p. 17 (type torquatus, presumably
from South America).

248 The University Science Bulletin

One female specimen (No. 4560) was collected at night along the
railroad about a kilometer from Tonala, Chiapas.

Dorsal scales in 17 rows ; ventrals, 197 ; tail broken ; third and
fourth supraoculars enter eye on one side, third only on the other;
6-7 supralabials; 7-7 infralabials; no preocular; two postoculars;
temporals, 1-2-3; two or three scales between single pair of chin-
shields and first enlarged ventral; one loreal; anal divided, preced-
ing scale entire; 262 mm. snout to vent.

Dorsal color dark tan, reaching to second scale row; a faint, nar-
row, median dorsal line of darker brown; belly, cream.

E. sumichrasti differs from the specimens referred to unicolor in
the character of the rostral and in the shape of the frontal and pre-
frontals. In unicolor the rostral is but very slightly produced, and
somewhat narrower as seen from the ventral surface. The lateral
edge of the rostral is anterior to the suture between the second and
third infralabial in unicolor, about even with the middle of the third
infralabial in sumichrasti. In the latter species the rostral is dis-
tinctly produced anteriorly, projecting beyond the mental a distance
slightly less than that between the posterior median edge of the chin-
shields and the median labial border of the mental. In unicolor the
length of the rostral on the median ventral line is equal to the dis-
tance between the posterior median edge of the first infralabials and
the median labial border of the mental. In the latter species, the
angle between the anterior and lateral edges of the prefrontals is
about 90 degrees; in sumichrasti the angle is considerably greater,
and as a result the prefrontals are not so wide at the side as in
unicolor. In sumichrasti the anterior edge of the frontal is slightly
convex, while in unicolor it is strongly convex. The frontal is pro-
portionately shorter in sumichrasti, its length not equaling its dis-
tance from the tip of the snout.

In unicolor the snout, in lateral profile, is distinctly down-curved,
while in sumichrasti it more nearly approaches a straight line.

Fischer's description {loc. cit.) of unicolor is inadequate for cer-
tain identification of his species; however, the figures accompanying
the description, while poor, show in general the features character-
istic of the specimens here referred to unicolor.

E. sumichrasti was described by Bocourt as having a preocular,
which character served as the basis for separation from murinus
Bocourt {longicaudatus Cope, a substitute name for murinus Bo-
court, which was stated to lack grooved teeth, present in m^urinus
Cope according to Cope). However, it appears that sumichrasti
and longicaudatus are synonymous, as the presence or absence of a

Taylor and Smith : Mexican Snakes

249

preocular is a variable character according to Dr. E. R. Dunn (in
litt.) , who states that it occurs on one or both sides of specimens
from Mexico, Panama and South America. E. sumichrasti has
priority over E. langicaudatus.

That sumichrasti is distinct from murinus Cope (type locality
Chinandega, Nicaragua) is indicated by information from Doctor
Dunn {in litt.), who states that the two cotypes of murinus, and the
one other existing specimen from Nicaragua, have a light nuchal
collar, lacking in both sumichrasti and unicolor.

Storeria dekayi (Holbrook)

Three, specimens (Nos. 4662-4), all from a locality five miles
south of Valles, San Luis Potosi. All were found in piles of drift-
wood near a river bank.

Third and fourth supralabials enter eye; supralabials 7-7 in
two, 6-7 in one (sixth and seventh supralabials fused on one side) ;
infralabials 7-7; postoculars 2-2 in two, 3-4 in one; one preocular;
loreal absent; anal divided; no scales between chinshields and first
enlarged ventral.

Measurements (in mm.) and scale counts of Storeria dekayi

Number.

Sex.

Ventral.s.

Caudals.

Total
length.

Tail
length.

4663 . . .

9

141
137
141

49
57
59

355
288
311

70

4664

70

4662

72

Storeria storerioides (Cope)

Ten specimens, from the following localities: Tres Cumbres
(Tres Marias), Morelos (No. 4665); Zempoala Lake, near Tres
Cumbres, Morelos (Nos. 5350-1) ; Desierto de los Leones, Distrito
Federal (Nos. 5354-5); Rio Frio, Puebla (No. 5403); 57 kilo-
meters southeast of Mexico City, on road to Puebla (No. 5352) ;
66 kilometers southeast of Mexico City, on road to Puebla (No.
5353) ; two miles south of San Martin, Mexico, near Zitacuaro,
Michoacan (No. 5404). One specimen lacks locality data. All
were found at elevations between 9,000 and 10,000 feet above sea
level, in pine forests. One was found under a log, the remainder
crawling about on the ground.

250

The University Science Bulletin

Fig. 3. Storeria storerioides (Cope) EHT-HMS, No. 5404; near Zitacnaro,

Michoacan, in Mexico. X5.

Scale rows 15-15-15; third and fourth supralabials entering eye,
except on one side in one, in which the third only enters the eye;
supralabials 7-7 in six, 6-6 in two, 5-7 in one, 7-8 in one; three
preoculars, on one side in one, two in others; postoculars, 2-2 in
eight, 2-3 in one, 3-3 in one; temporals, 1-1-1 in one (sides counted
separately), 1-2-1 in one, 1-2-2 in ten, 1-3-2 in one, 1-2-3 in six,
1-3-3 in one; one loreal, entering orbit between preoculars on one
side in one, separated from orbit in others; anal divided; second,
third and fourth subcaudals entire in one specimen, remainder
divided. The nasal is divided below the naris in all specimens, and
is divided above the naris (apparently) in three. In eight speci-
mens the angle formed by the two posterior sides of the frontal is
90 degrees or less, and the frontal extends posterior to a line be-
tween the posterior edges of the supraoculars. In two specimens
the angle formed by the two posterior edges of the frontal is about
110 degrees, and the frontal extends posteriorly only to a line be-
tween the posterior edges of the supraoculars.

One specimen, the smallest examined (No. 5404), has the scales
of the lateral row of dorsals keeled. This specimen exhibits several
other variations, such as a divided nasal, a short, broad frontal
with an obtuse posterior angle, and the first enlarged ventral in
contact with the posterior pair of chinshields (separated by one
scale in others), some of which may be due to its immaturity.

In coloration, the specimens agree with descriptions published.

Taylor and Smith: Mexican Snakes

251

It may be added that an irregular, narrow, reddish-brown band
extends the length of the body on the middorsal line.

Measurements (in mm.) and scale counts of Storeria storerioides

Number.

Sex.

Ventrals.

Caudals.

Total
length.

Tail
length.

4665

9

131

47

127

26.5

5350

9

45

236

47.0

5354

9

132

46

255

50.0

5403

9

129

46

259

52 0

5404

cf

131

51

110

23.7

5352

&

130

49

137

29.5

5353

cf

129

50

140

31.0

5356

d'

130

50

172

39.0

5351

cT

131
130

53

51

222

288

51.5

5355

64.0

Chersodromus liebmanni Reinhardt

Chersodromtis Liebmanni Reinhardt, Vid. Meddel. Naturh. Foren. Kjobenhavn, 1860, pp.
243-245, Plate IV, figs. 10, 11 (type description; type locality, Mexico).

A single specimen of this species EHT-HMS, No. 4564, was
collected in a coffee plantation, 10 mi. E. of Cordova, Veracruz,
July 23, 1932 (Taylor), from under a small rotten log. From under
a fallen banana stem contiguous to this log was captured at the same
time a specimen of Ninia diademata Baird and Girard. These two
snakes bear a very strong superficial resemblance to each other.
Both are black with yellow collars, their bodies nearly of the same
thickness. Actually they are very different.

The following data are taken from No. 4564: Ventrals, 135; anal,
entire; subcaudals, 34; supralabials, 7-7; lower labials, 8-8; pre-
ocular, 0-0; loreal, 1-1; postocular, fused with supraocular; tem-
porals, 1-2; total length, 199; tail, 33.

Supranasals wider than long; prefrontals completely fused; frontal
more or less triangular, as wide as long, as long as its distance
from tip of snout, shorter than the parietals; nasal divided nearly
equally, the nostril lying between; rostral barely visible above;
five labials touch the very large anterior chinshields which are very
much larger than posterior; the first labials not in contact; scale
formula, 18-17-17-17. Scales are keeled dorsally, dimly anteriorly,

252 The University Science Bulletin

then becoming stronger and more evident posteriorly; the outer
scale row much enlarged.

The type has 130 ventrals ; subcaudals, 42 ; its length is 250 mm.

Ninia diademata Baird and Girard

Ninia diadeinata Baird and Girard, Catalogue of North America Reptiles, Part I, Serpents,
1853, p. 49. (Type description; type locality, Orizaba, Mex. Jas. Fairie coll.)

A single specimen (EHT-HMS No. 4565 $ ) was collected in a
coffee plantation 10 mi. E. of Cordova, Veracruz, July 23, 1932
(Taylor), from under a banana stem, as mentioned in the discussion
of Chersodromus liehmanni.

The following characters are evident: Ventrals, 143; subcaudals,
96; upper labials, 6-6; lower labials, 6-6; preoculars, 0-0; post-
oculars, 2-2; temporals, 1 + 2; scales touch chinshields, 4-4; anal,
undivided; total length, 268 mm., tail, 92 mm. Rostral barely visible
above; internasals about one third size of the prefrontals; latter
large, entering the eye above the loreal; frontal hexagonal, as wide
as long, minutely longer than its distance to tip of snout; parietals
longer than frontal, equal to or a little greater than their distance
from snout tip; nasal divided; loreal rectangular, entering the eye;
first chinshields double size of second pair; scales very heavily
keeled and striated, the formula 21-19-19-19. Outer row rather
large; the others become smaller towards middle of back.

This specimen differs from the type in the absence of preoculars,
and in having the prefrontals enter the orbit. It is presumed that
the species is variable in these characters. So far as I can find other
specimens referred to this form do not agree with the type.

The coloration is typical; the black median ventral line occupies
as much area as the cream lines bordering it. Outer edges of the
ventrals black.

Conophis vittatus Peters.

(Plate XXIII, fig. 1)

Conophis vittatus Peters, Monatsb. Akad. Wiss. Berlin, Oct., 1860, pp. 519-520; PI.,
fig. 3; (type description; type locality uncertain): "Diese Schlange wurde bei einem
Handler in Hamburg gekauft, welcher mir erzahlte, dass sie nicht weit von Neu-Orleans in
Mississippi mit einem treibenden Strohhausen auf ein Schiff gezogen sei."

Four specimens were taken; EHT-HMS Nos. 5149, 5150, 12
mi. S. Puente de Ixtla, Morelos, km. 133, July 14, 1936, and Aug.
1, 1936; No. 5152, km. 350, between Rincon and Cajones, Guer-
rero, July 24, 1936; No. 5151 Xaltianguis, Guerrero, July 27, 1936
(km. 405) ; Taylor collector.

The color pattern in the specimen figured is also that of the

Taylor and Smith: Mexican Snakes 253

other three specimens. The ground color is light olive or yellowish-
brown. Data here given arc from Nos. 5149 5 ; 5150 $ ; 5151 ^ ;
5152 ^ , respectively:

Scale formula, 29, 19, 19, 17; 28, 19, 17, 17; 26, 19, 17, 17; 29,
19, 19, 17; ventrals, 165, 163, 162, 160; subcaudals, 48, 59, 70, 67;
upper labials, all 7-7; lower labials, 9-9, 10-9, 9-9, 9-9; chinshields
of equal length; four scales touch chinshields. Measurements in
millimeters: Total length, 500, 561, 522, 552; tails, 77, 110, 133,
117; head width, 13, 12.4, 13, 13; head length, 14.2, 20, 20, 19.

One of us (Taylor) was bitten by No. 5151 on the middle joint
of the third finger. Pain and swelling ensued which lasted for some
time. A year and a half after the occurrence finds the digit still
somewhat weak, and slightly painful when bent.

Conophis concolor Cope

Conophis concolor Cope, Proc. Acad. Sci. Philadelphia 1866, p. 318 (type description; type
locality, Mexico).

H. M. Smith obtained a single specimen at Chichen Itza, Aug.
26, 1936 (EHT-HMS No. 11635). It differs from the type descrip-
tion in being rather olive brown. Paired dotted lines appear on
back of head and for about an inch on the neck. The black stripe
through the eye continues as a dim, dotted line for an inch on the
neck; a dim, lighter line is suggested between first and second scale
rows. Chin dark with yellow spots on posterior lower labials;
upper labials with a cream line. Further data on this specimen is
given elsewhere (Smith, this journal).

Tantilla rubra Cope

Tantilla rubra Cope, Jouni. Acad. Nat. Sci. Philadelphia (2) VIII, 1876, p. 144 (type
description; type locality, Japana, Tehuanfepcc, Dr. Sumichrast coll.).

This species is represented by three specimens, EHT-HMS Nos.
5240 s (Radycliffe Roberts) ; 5241 $ , 5242 ^ (Taylor) from Km.
226, 22 km. N. W. of Tehuacan Puebla. Scale formula, 16-15-15,
15-15-15, 15-15-15; ventrals, 154, 156, 160; subcaudals. 66, 62, 60;
upper labials, all 7-7; lower labials, all 6-6; preoculars, all 1-1;
postoculars, all 2-2; temporals, all 1 -f (the second lower temporal
is behind last labial).

Measurements (in riim.) : Total length, 348, 320, 231; tail, 88,
80, 54; head length, 11.4, 10, 9; head width, 6.5, 6, 5.7.

Superficially this species bears a rather close resemblance to
Tantilla bocourti, but the details of the head coloration differ. The
head, likewise, is black generally; the black and yellow nuchal

254 The University Science Bulletin

collars are much the same, save that the yellow collar encroaches
more on the parietals; behind the eye is a more or less rounded
spot of yellow, the lower border of which is formed by the black
spots on the lower labials; tip of snout yellowish, with small black
spots in front of nostrils and one on rostral; sometimes small spots
on the outer part of the internasals.

In life the color is a light shade of brownish-red, becoming faun
color in alcohol. These specimens were obtained from under cactus
plants. Minute pits, comparable to apical pits of other genera, are
present on the scales of this species back from the tip of the scales
near the middle. These cannot be seen unless a strong lens is used.

Tantilla bocourti (Giinther)

Homalocranium bocourti Giinther, Biologia Centrali Americana; Reptiles, Batrachians, Jan.,
1895, p. 149; (type description; type locality, "Guanajuato."); Taylor. Trans. Kansas
Academy of Science, 39, 1936, pp. 336-337, fig. 1.

Five specimens are at hand: EHT-HMS, No. 5239 ^ Cuerna-
vaca, Morelos; Taylor, July 12, 1936, Nos. 2245 $ , 2244 s , ^^
km. 226, 22 km.; N. W. Teliuacan, Puebla; Taylor, Aug. 23, 1936,
Nos. 14430 ^ , 14431 5 , Cuernavaca, Morelos; H. M. Smith, Oct.
17, 1936.

The ventral and subcaudal counts of these specimens are, respec-
tively, 174, 57; 173, 46; 164, 58; 177, 54; 185, 47. The specimens
from Cuernavaca are generally darker than those from Tehuacan,
and with a higher ventral-subcaudal count. From data here given,
and published data, the following ventral-subcaudal counts obtain.
The type has a total count of 227 (Guanajuato) ; Magdalena,
Jalisco, 228-238; Cuernavaca Morelos, 231-234; Tehuacan, Puebla,
Acultzingo, Veracruz, 219-221. An east-west range of from 238
to 219 is discernible.

Tantilla calamarina Cope

Tantilla calaviarina Cope, Proc. Acad. Nat. Sci. Philadelphia, 1866, p. 320; (type descrip-
tion; type locahty, Guadalajara). Taylor, Trans. Kansas Acad. Sci. 39, 1936, pp. 346-347,

fig. 5.

A specimen collected four miles S. Cuernavaca (EHT-HMS No.
5243 5 ) has a broader head than the specimen from Queseria,
Colima, figured by Taylor, loc. cit. Ventrals, 130; subcaudals, 27;
upper labials, 6-6; lower labials, 6-6; preoculars, 1-1; postoculars,
1-1. Temporals, 1 + 2; mental touches chinshields; temporal in
contact on both sides with the postocular. Total length, 202 mm. ;
tail, 27 mm.

Taylor and Smith : Mexican Snakes 255

Three dim, blackish stripes, one median, the outer on third and
fourth scale rows, the space between forming slightly lighter lines.

Stenorhina degenhardti quinquelineata (Hallowell).

(Plate XXIII, fig. 2)
Microphis quinquelineata Hallowell, Proc. Aratl. Nat. Sci. Philadelphia, 1854, p. 97;
(type description; type locality, Honduras).

A specimen (EHT-HMS, No. 4567 2 , Totolapam, Oaxaca, Aug. 6,
1935, H. M. Smith) of a form agreeing in detail with the characters
given (and figured) for quinquelineata by Hallowell, causes me to
resurrect Hallowell's name. It differs from fremenvillei in having
a broad median black line, and two narrow, lateral black lines on
each side; the ground color between the dark lines is a putty-gray,
the edges of the scales slightly darker. Between the median and
upper lateral lines is an indistinct dotted, dark line; below uniform
yellow. The lips are bright yellow, without spotting.

Compared with the form apiata this snake has a shorter snout,
the prefrontals and internasals distinctly shorter; the snout is ac-
tually broader; and the frontal is narrower and more elongate.

The specimen presents the following characters: Ventrals, 175;
caudals, 34; scale formula, 24, 17, 17, 17; upper labials, 7-7; lower
labials, 7-7; 3 labials touch anterior chinshields; one labial touches
second chinshields; 3d and 4th labials enter eye; part of rostral
visible above, equal to three fourths its distance from the frontal;
latter elongate a third longer than its distance from the end of the
snout; outer anterior corner of the parietals segmented, forming a
second anterior "temporal"; diameter of eye equals the distance to
nostril (in apiata much less) ; nasal segmented, the anterior moiety
fused with the internasals; loreal wanting; preocular touching pos-
terior nasal; 1 preocular, two postoculars; temporals, 2-2.

Total length, 465 mm.; tail, 60 mm.; head width, 11mm.; head
length, 15 mm.

Stenorhina degenhardti apiata Cope

Stenorhina degenhardti apiata Cope, Journal Acad. Nat. Sci. Philadelphia, (2) VIII, p. 142;
(type description; type locality, Tehuantepec) Sumichrast, collector.

This form is represented by a specimen (EHT-HMS, No. 5153)
collected five miles north of Acapulco at Puerto Crucita. The speci-
men was bright pinkish-red above; the ventral surface of a similar
color, but of a lighter shade. There is an indistinct dark line be-
hind eye, and a trace of a median dorsal line on the neck. In
alcohol the snake has become light brown, and where the scales are

256

The University Science Bulletin

lost it is nearly flesh color. The upper labials are bright yellow,
the chin and lower labials creamy-yellow. Scale formula, 24, 17,
17, 17.

The accompanying figure shows scale relationships. Ventrals,
172; subcaudals, 32; anal divided; total length, 490mm.; tail,
65 mm.; head width, 10 mm.; length, 15.2 mm.

A

Fig. 4. Stenorhina degenhardti apinla (Cope). EHT-HMS, No. 5153;
five miles north of Acapulco. Actual head width, 10 mm.

Taylor and Smith: Mexican Snakes

257

1 ji •

y.

^'.J

PLATE XXIII

Fig. 1. Curiopln.s rittdtus Peters. EHT-HMS, Xo. 5149 9 ; total length
500 mm. '

Fig. 2. Stciiurlnna dcyoihardti quinqucliiudla (Hallowell) EHT-HMS No
4567 9 ; total length, 465 mm.

^ Fig. 3. ('inii,j>.^i.s jrontalk (Cope). EHT-HMS, Xo. 5203 9; total length
570 mm. '

Fig. 4. XoioiUm anguslirosim Peters. EHT-HMS, No. 5207 9- total
length, 587 mm. '

17—4141

258 The University Science Bulletin

Clelia clelia (Daiidin)

Coluber clelia Daudin, Hist. Nat. Gen. Paitif. Rfiit. vol. 6, year XI (1803). pp. 330-331,
pi. LXXVIII.

A specimen (EHT-HMS), No. 4568 ^ , was collected by H. M.
Smith at Paso del Rio, Colima. Ventrals, 211; caudals, 86; upper
labials, 7-7; lower labials, 8-8; preoculars, 1-1; postoculars, 2-2;
temporals, 2 + 3; anal single; scale formula, 24, 21, 17, 17, 17;
preocular separated from frontal; parietals shorter than their dis-
tance to end of snout; frontal about equal to its distance to end
of snout. Eye (3 mm.) shorter than distance between eye and
nostril (3.6 mm.); loreal higher than long; length, 675 mm.; tail,
147 mm. ; head width, 12 mm. ; length, 19.6 mm. Head dark violet to
violet-black; cream collar behind parietals 4 scale-rows wide; large
blackish violet band behind collar.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV 1 June 1, 1938 [No. 14

Concerning Mexican Salamanders

EDWARD H. TAYLOR,

DepartiiR'nt uf Zoulog.w University of Kansas

Ab.stract : This study is based on a large collection of Caudata made in
Mexico during the years 1932-1936, chiefly by Edward H. Taylor and Hobart
M. Smith. Three species are described from the collection of the Museum
of Comparative Zoology, Harvard College.

Oedipus robertsi (Nevada de Toluca, Mexico), Oedipus smithi (Oaxaca),
Oedipus altamontanus (Morelos), Oedipus giganteus (Jalapa, Veracruz),
Oedipus multidentata (San Luis Potosi). Oedipus matnii (Hidalgo), and
Amhystomn schmidti (Mexico) are described as new. Oedipus leprosus (Cope),
recently placed in the synonymy of Oedipus cephcdicus (Cope), is revived as
a valid species. The following species are discussed : Oedipus bellii, chiropterus,
cephalicus, leprosus, orizahensis, platydaclylus, salvinii, lineolus, penuntulns;
Ryacosiredon altamirani; and Gymnopis multiplicata oaxacae.

THE present study is based on the collections made by Hobart
M. Smith and myself in various localities in central and south-
ern Mexico. Several of the species are represented by large series;
some, unfortunately, by only one or two specimens or at most a
small scries.

Dunn, in his admirable work on the Salamanders of the family
Plethodontidae, (1926) deals extensively with the plethodontid
genus Oedipus, uniting with it the distinctive forms sometimes rec-
ognized under the generic designations Oedipus Keferstein {Ophi-
obatrachus Gray) and Thorius Cope. This work has been especially
helpful in the study of the genus Oedipus.

The recent review of the Ambystomid Salamanders of Mexico by
Lafrentz and Wolterstorff (Abh. Ber. Mus. Natur. — Heimatk.
Natur. Ver. Magdeburg, Bd. VI, Heft. H, pp. 90-127 and 128-149,
respectively), has been helpful in the study of Ambystoma and
Ryacosiredon.

I am under great obligation to Dr. Thomas Barbour and Mr.
Loveridge for the loan of the salamanders of the genus Oedipus in

(259)

260 The University Science Bulletin

the Harvard collection and for the privilege of describing three new
forms.

The following recognized species are known to occur in JNIexico
(elevation, in feet, is an approximation):

Family PJ( Ihndontida

c

Oedipus belUi (Gray) lowland (?) to 8.000

Oedipus giganteus sp. nov 6,000 to 10,000

Oedipus gadovii Dunn 6,000 to 14,000

Oedipus sulcatus (Brocchi) 5.000

Oedipus chiropterus (Cope) 4,000 ( ?) to 11,000

Oedipus multidcntaLus sp. nov ? 8,000

Oedipus rohertsi sp. nov 10.000

Oedipus altcmiontanus sp. nov 10,000

Oediptis smithi sp. nov 9,000 to 10,000

Oedipus macrinii Laf rentz 3.270

Oeelipus manni sp. nov 8,000

Oedipus cepholicus (Cope) 8.000 to 13,000

Oedipus orizabensis (Blatchley) 8,000 to 12,000

Oedipfus leprosus (Cope) 10,000

Oedip)is platydactylus (Ciivier) lowland

Oedipus salvinii (Gray) lowland

Oedipus yucatanus Peters lowland

Oedipus rufescens (Cope) lowland

Oedipus lineolus (Cope) lowland to 4,000

Oedipus towrisendi Dunn 4,400-? 8,000

Oedipus petvintidus (Cope) 8.000 to 10,000

Batrachoseps attenuatus leucopus Dunn lowland

Ensatina eroecnter (Cope) 3,600 to 7,000

Family Amhystomidae

Avihystoma schmidii sp. nov about 8.000 feet

Ambystoma mexiccnntm Shaw about 8,000 feet

Ambystoma dumereli Duges /?

Ambystoma tigrinum velascoi Wolterstorff . . . . 6.500 to 7.500 feet

Ainbystoma tigrinum tigrinum Green* lowland

Rlu/arosiredon altnniirani Duges about 11,000 feet

Family Scdamandrielae

Triturus meridionalis Cope lowland

Triturus kallerti (Wolterstorff) lowland

Triturus torosus Eschscholtz lowland

Order Apoda

Derm,ophis mexicanus Dumeril and Bibron lowland

Gymnrtpis multiplicata oaxaeae Mertens lowland

It is to be regretted that more detailed data on vertical distri-
bution was not obtained. The value of such data is evidenced by
the recent study on salamanders of Guatemala, by Schmidt (1936).

Several species names proposed or used for Mexican forms have
been regarded as synonyms of previously described species.

* Aiiihystoma prosrrpiiie Biiird and Girard. Proc. Acad. Nat. Sci. Phila. 1852, p. 173.

Taylor: Mexican Salamanders 261

1. Spelerpes orcidus Cope. This species has long been regarded
as a synomym of Oedipits chiropterus (Cope), having been placed
in the synonymy of that species by Cope, himself (1869). I have
not examined the type.

2. Bolitoglossa mexicana Dumeril and Bibron (1854), in part.
(Erp. Gen. Atlas, plate 104, fig. 2.) is referable to Oedipus bellii.

3. Spelerpes minimus Wiedersheim. Dunn (1926) , declaring it a
nomen nudum, settles this name in the synonymy of Oedipus pen-
natulus (Cope).

4. Spelei^pes mono Boulenger (1882), Jalapa, Veracruz (non
Cope). Probably based on more than one species, but which I can-
not say.

5. Spelerpes laticeps Brocchi (1883). The description is brief,
and I am uncertain whether this species should be recognized or
not. The type should be examined and redescribed. The brown
color, as shown in the figure (Brocchi, p. XVHI, fig. 1), may be due
to the preservation.

6. Geotrition carbonarius Cope (1860), Jalapa, Mexico, is ap-
parently referable to Oedipus platijdactylus. Boulenger (1882)
places it in the synonymy of Spelerpes variegatus = [Oedipus
platydactylus (Cuvier)].

7. Spelerpes punctahmi Brocchi (1883). This has been placed in
the synonymy of Spelerpes variegatus Gray = {Oedipus platy-
dactylus], but the type should be reexamined.

8. Spelerpes attitlanensis Brocchi (1883). Schmidt (1936), who
examined the types, regards this a synonym of Oedipus salvinii
(Gray).

9. Spelerpes gibbicaudus Blatchley (1893) Mt. Orizaba, Veracruz.
This species is founded on the type specimen of Spelerpes leprosus,
and is a synonym of leprosus.

Rhyacosiredon Dunn

1928. Rhyacosiredon Dunn, Proc. New England Zool. Club, Vol. X, Nov. 3, 1928, pp.
85, 86.

1930. Amhystortia Lafrentz, Abb. Ber. Mus. Natur-Hcimatk. Naturw. ver. Magdeburg,
Bd. VI, Heft. 2, 1930, p. 115.

1930. Ambystoma (Rhyacosiredon) Wolterstorff Abh. Ber. Mus. Natur-Heimatk. Naturw.
Ver. Magdeburg. Bd. VI, Heft. 2, 1930, p. 142.

Dunn characterizes the genus as follows: "An Ambystomid
salamander with no gills or gill slits in the adult state; lungs pres-
ent; ypsiloid well developed; no free lachrymal; nasal present;
adult with vomerine teeth in larval position; maxilla very small;
premaxillary teeth aborted, not as long as bony edge of premaxilla;

262 The University Science Bulletin

both jaws with horny beak in adult and larva; larvae with gills
with long rami; dorsal fin aborted in large larvae; eggs (ovarian)
larger than those of other species, 3 mm. in diameter."

Bhyaco sire don altamirani (Diiges)

1895. Avibystoma altamirani Dug?.s, Description d'un Axolotyl de Montagnrs de las
Cruces (Anibystoma altamirani. A. Duges), 1895, No. 15. Imprimerie du Ministere de
"Fomento" (Type description; type locality "Manantia! de los Axolotes en la Serrania de las
Cruces, perteneciente al Valle de Mexico"); and La Naturaleza (2), II, 189*i. pp. 459-4(il,
pi. XIX; Lafrentz, Abh. Ber. Mus. Natur-Heimatk. Naturw. Ver. Magdeburg, Bd. VI,
Heft. 2, 1930, pp. 115-120 (numerous localities in mountains near Mexico City).

1928. Rhyacosiredon altamirani Dunn, Proc. New England Zool. Club, Vol. X, Nov. 3,
1928, pp. 85, 86 (Santa Fe, Contreras 8,090 feet, Dos Rios 8,800 feet, all in the Ajusco
Moimtains south and west of Mexico City).

1930. Ambystoma (Rhyacosiredoyv) altamirani Wolterstorff, Abh. Ber. Mus. Natur-
Heimatk. Naturw. Ver. Magdeburg, Bd. VI, Heft. 2, 1930, pp. 142-144, fig. 11.

Two specimens (EHT-HMS, Nos. 12511, 12512) in the collection
were captured under logs, near a small stream flowing into Lake
Zempoala, at an elevation of about 11,000 feet.

While agreeing in most characters, these two specimens differ in
certain points which may be due to the age of the specimens.

No. 12511. Body with numerous distinct black spots on the
dorsal and dorsolateral surfaces; head spotted likewise. Tail
clouded with lighter and darker mottling; many maxillary teeth
seem to be missing, there being only about six on each side, these
covered with fleshy gums; six premaxillary teeth, elongate, curving.
Vomerine teeth in elongate series which converge, but fail to meet
anteriorly by a distance equal to more than double the diameter of
the rather large choanae; each series is broken; the posterior group
of six teeth is slightly curved around the posterior part of the
choanae to a point nearly opposite the middle of choanae, where
a break, equal to the width of a choana, occurs; the following eleven
teeth form a straight continuous series.

No. 12512. Color, generally drab olive, the dark spotting not
or scarcely discernible; the tail is mottled, similar to the previous
specimen. Maxillary and premaxillary teeth form an unbroken
series of 17 (18) teeth in each half of the jaw; the premaxillary
series consists of 8 teeth.

In both specimens the limbs are very large and the body very
short. Limbs when adpressed overlap the length of the arm from
elbow. Metacarpal and metatarsal tubercles very distinct, large,
rather close together. Twelve costal grooves; no groove behind
eye; interorbital distance very wide; a skinfold on lower lip; tips
of digits brownish ; no enlarged mucous pores on head.

Taylor: Mexican Salamanders 263

Measi(rc77U'nts (in mm. I of Xus. 12511, 12512, respectively: Snout
to vent, 68.2, 64.6; snout to arm, 25, 23.3; tail, 75, 68.5; width of
head, 18, 16.5; length of head, 22, 19; axilla to groin, 33.5, 32;
foreleg, 27, 24.5; hindleg, 29, 25; interorbital distance, 5.3, 5.4; eye
to nostril, 4.4, 3.5; length of snout, 5.2, 4.4; eyelid, 2, 2; eye, length,
3, 3.2; between nostrils, 5.1, 5.

I believe that Dunn is fully justified in placing this form in a
genus separate from Amby stoma.

It will be noted that these specimens display nothing that can be
construed as horny beaks. There is, however, a slight deposition of
keratin along the borders of the lips, which is scarcely noticeable.
This condition may be due to the age of these specimens.

Ambystomn schmidti sp. nov.

Holotype. EHT-HMS No. 3999, collected 10 miles east of San
Martin (Asuncion) at Rancho Guadalupe Aug. 3, 1932. E. H.
Taylor, collector.

Diagnosis. A very small species of Atnbystoma, with short limbs
and relatively small feet ; adpressed limbs separated by three costal
folds; 14 costal grooves (an axillary fold apparently wanting);
length of eye about equal to the distance of the eye from the nostril;
width of eyelid contained more than two and one half times in the
interorbital distance; a nuchal fold; tail shorter than the head and
body. Tongue with a deep median groove; series of vomerine teeth,
beginning behind choanae near their posterior inner border, form an
arch between choanae; two tubercles on hand and foot; four pha-
langes in fourth toe.

Description of the type. Head flat with trace of a canthus
rostralis; nostrils nearer tip of snout than eye, the distance between
them about four fifths of interorbital distance; a slight median oc-
cipital depression with very slight longitudinal swellings; a slight
median dorsal groove along the middle of the back; a well-defined
fold across underside of neck, continued as a groove on the side of
the neck, but not continued across the dorsal surface; a groove be-
ginning in the upper temporal region crosses the angle of the jaw
and passes beneath the chin to meet its fellow from opposite side; a
deep groove from eye runs back to the lateral nuchal groove, where it
terminates; below the posterior part of this groove is a fold of skin
terminating at the lateral nuchal groove; 14 costal folds (that in
axilla apparently wanting); 12 grooves cross the abdomen; lips of
anal slit swollen, the walls showing some slight foldings; limbs
rather short, separated by three costal folds when adpressed; first

264 The Uxiversity Science Bulletin

finger well developed, the second and third largest, of nearly same
size; fourth toe much longer than first; all digits flattened some-
what with a slight skinfold along their edges; a slight indication of
a web between toes; foot broad, the ascending order of length of toes,
1, 5, 2, 3, 4; toes rather bluntly pointed; tail compressed, with only a
slight trace of a crest for a short distance on the base of the tail.
Skin smooth, the head showing minute pitting.

Vomerine teeth in a continuous series consisting of about 16 teeth ;
they begin behind the choanae near the inner posterior edges and
curving between choanae, fail to reach the level of their anterior
edge; a median depression in the palate anterior to the vomerine
teeth; groove from the choanae covered by a triangular projection
from side of jaw; maxillary and premaxillary teeth about forty on
each side; a few other scattered teeth behind the regular series, an-
teriorly.

Color. Above violet to lavender with small scattered cream spots
on sides of head, body and tail; tail somewhat darkened on side; be-
low yellow-cream, the color extending somewhat on sides; upper
side of limbs somewhat lighter than body.

Measurements (in mm.). Snout to vent, 52; tail, 37; head length,
10; head width, 9.3; snout to arm, 15; axilla to groin, 28.4; arm, 12;
hand and finger, 4.3; leg, 14; foot and longest toe, 7.5; eye length,
2.3; interorbital distance, 3.5; eyelid, 1.25.

Remarks. Tlie specimen was obtained from under a log in a pine
forest near a large artificial pond. It is not, apparently, closely re-
lated to the other recognized Mexican species of the genus, as evi-
denced by the absence of large pits on the head, and the very small
limbs. The elevation of this locality is probably 8,000 feet.

This species is named in honor of Dr. Karl D. Schmidt, of the
Field Museum, Chicago, who has had the kindness to furnish me
with comparative salamander material from Central America.

Oedipm belUi (Gray)

(Plate XXVII; figs. 1, 2)

1849. Oedipus platydactylus Baiid, Journ. .^cad. Nat. Sci. Philadelphia, (2), 1, 1849, p.
286 (Not Salamaiidra platydactyhis Cuvier).

1850. Spelerpes bellii Gray, Cat. Batr. grad. Brit. Mus.. p. 40 (type description; type
locality. "Mexico"); Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 372; and 1869, p.
105; and Proc. Auier. Philos. Soc, 18, 1879. p. 263; and idem, 1869, p. 105; Boulenger,
Cat. Batr. grad. Caud. British Mus., (2), 1882, pp. 68-69; Brocchi, Mission Scientifique
au Mexi(|U' et dans I'Anierique Centrale, Etude sur les Batraciens, Liv-r. 3, 1883, p. 110, pi.
206 (part.); Ferrari-Perez, Proc. U. S. Nat. Mus., 9, 1886, p. 199; Cope, Bull. U. S. Nat.
Mus., No. 32, 1887, p. 8; and Bull. U. S. Nat. Mus., No. 34, 1889, p. 161; Blatchley,
Proc. U. S. Nat. Mus., 16, 1895, p. 37 (Part.); Duges, La Naturaleza, (2), 2, 1896, p.
482; Moore, Proc. Acad. Nat. Sci. Philadelphia, 1900, p. 019: Ciiinther. Biologia Centrali-

Taylor: Mexican Salamanders 265

Americana, Rept. and Batr., 1902, 11. 299 ; Dr Li'uu, Iiulice di' lus Batiafios que se
encuentran en la Repiiblica Mexicana, June, 1904, p. 37; Gadow, Zool. Jahrb., 1910, pp.
709, 714.

1854. Bolitoglossa Mexicana Dunioril and Bibon, Erp. Gen, 9, 11. 93. pi. 104, fig. 2
(Oaxaca, Mexico; Veracruz, Mexico); Duges, La Naturaleza, I, 1869, p. 144.

1854. Salaniandra togata Valenciennes, mentioned in Dumeril and Bibron, Erp. Gen. 9,
1854, p. 94 (apparently not published).

1856. Spelerpes Mexicana HallowoU, Proc. Acad. Nat. Sci. Philadelphia, 1856, ji. 11.

188 4. Geotriton belUi Garman, Bull. Essex Inet., 16, 1884, p. 471.

1918. Oedipus beliii Dunn. Bull. Mus. Comp. Zool., 1918, 62, p. 471 ; Field Mus. Nat.
Hist. Zool. Ser, XII, pp. 99, 100; The Salamanders of the family Plethodontidae, Smith
College 50th aniv. publ., 1926, pp. 357-360, fig. 57, map (part.); Wolterstorff, Abh. Ber.
Mus. Nat. Heimatk. Naturw. Ver. Magdeburg, band, VI, Heft. 2, 1930, p. 140.

This, one of the most striking species of the Mexican salamander
fauna, enjoys a wide distribution on the plateau region, but ap-
parently occurs sporadically. A series of specimens (Nos. 3973-
3994) in the collection was taken by Hobart M. Smith and David
Dunkle near Belen, 40 kilometers north of Teocaltiche, Jalisco, July
22, 1934.

Description of the species. (From EHT-HMS. No. 3981 ^ .)
Large species with robust body; seen from above the head is truncate
oval, flattened; eye large, longer than the snout, but about equal to
its distance from the tip; the posterior parts of eyelids inserted un-
der a fold; a groove from behind eye w^hich joins the first gular
grooves; latter short, not joining in the middle of throat; gular fold
present, prominent; 13 costal grooves; three costal folds between
adpressed limbs; limbs well developed, the toes, in descending
order of length, 3, 4, 2, 5, 1; both fingers and toes somewhat webbed
at base, flattened; tail longer than body, somewhat circular, with a
basal constriction; anal lips lined with papillae (male); vomerine
teeth in two elongate series, curving back strongly, medially, ex-
tending outward beyond the outer level of choanae in a straight line;
series narrowly separated medially ; parasphenoid teeth in two series
more or less contiguous anteriorly, but separated from each other
most of their length; separated from the vomerine series by a dis-
tance of little more than the width of a choana.

Color. Coal black on back and sides, somewhat grayish black
below; two large orange-yellow blotches on back of the head; a
somewhat inverted V-shaped spot on neck, followed by paired
series of orange spots extending down to and onto base of tail,
one pair corresponding to a costal fold.

Measurements of Oedipus beliii (Gray) (in mm.). Snout to vent,
93; snout to gular fold, 21; snout to foreleg, 29; axilla to groin, 53;
head width, 14.5; foreleg, 21.5; hind leg, 22; head width in head-
body length, 6.4 times; head length in head-body length, 4.6 times.

266 The University Science Bulletin

Remarks. A very young specimen (No. 3992), (19 mm. from snout
to vent) shows only a suggestion of the dorsal coloration, many of
the dorsal spots being missing; No. 3993 is practically the same.
No. 3984 (45 mm.) has many of the orange spots missing on the
right side, a few on the left side.

For the most part the specimens conform to the coloration and
marking of the specimen described. Specimens from Guerrero,
Hidalgo, have much thicker tails. The apparent differences may be
due to different methods of preservation.

Distribution. Known from Jalisco, Nayarit, Michoacan, Guerrero,
Oaxaca, Veracruz, Guanajuato, Queretaro, Hidalgo, and Distrito
Federal. Certain records of 0. bcllii for Veracruz are referable to
another species.

Larger series of this species will doubtless permit the separation of
certain subspecific groups in this widespread form. The record of
this species from Fort Whipple, Arizona, 3 specimens should be
questioned. (Listed by Dunn, 1926, and apparently doubted by
him.)

Oedipus giganteus sp. nov.

(Plate XXVI I. figs. 3, 4)

1883. Spelerpet! bellii Brocchi (part.). Mission Scientifique au Mexique et dans IWmerique
Centrale, Etude des Batiaciens. Livr. 3, 1883. pp. 110-11, ? pi. XX bis. fig. 2.

1826. Oedipus hellli Dinin (pari.). Salamanders of the Family Plethodontidae, Smith
Coll. Publ., 192G, pp. 357-360 (certainly MZC specimens Nos. 8434-8437, perhaps others).

Type. MCZ, No. 8435, Jalapa, Veracruz, Dr. E. R. Dunn, col-
lector ("under rocks, around the roots of trees in comparatively
open pastures").

Paratijpes. MCZ, Nos. 8434, 8436, 8437. Same collector and lo-
cality; EHT-HMS, 12040, 12085, Cofre de Perote, Veracruz, Elev.
10,000 feet.

Diagnosis. The largest species of the genus, related to Oedipus
belli, but differing in larger size, proportionally shorter axilla to
groin measurement, and proportionally greater head width; pits on
head well developed; large orange spots absent on occipital region;
adprcssed limbs separated by one fold or less in males, in the large
female by three folds; a median pit in roof of mouth between
choanac; vomerine teeth, 25-25; maxillary teeth, 47-45 in males; 4
premaxillary teeth pierce the lip in males; 68-72 maxillary-pre-
maxillary teeth in female; 62-62 mandibular teeth in males; para-
sphenoid series fused together anteriorly, diverging strongly pos-
teriorly.

Description of the type. Adult male. Head broader than neck;

Taylor: Mexican Salamanders 267

snout truncate, the nostrils widely separated; subnarial swellings
very moderate; interorbital region somewhat depressed, flattened;
eye (4.9 mm.) about equal to length of snout (4.8 mm.) ; smallest in-
terorbital distance (4.2 mm.) about equal to width of eyelid; width
between nostrils, 7.8 mm.; width of head (17 mm.) contained in
distance between snout and posterior part of vent (110 mm.), 6.4
times; head length (19 mm.) in same distance, 5.8 times; dorsal
surface of head and eyelids with well-developed pits, closely placed,
giving the skin a somewhat corrugated appearance; posterior corners
of eyelids fitting under a diagonal fold of skin; maxillary teeth, 47-
45; premaxillary teeth, four visible, piercing upper lip; mandibular
teeth, 62-62; vomerine teeth in two arched series of 25 teeth each,
separated medially by a distance less than width of choanae, ex-
tending much beyond choanae; parasphenoid teeth in two groups,
contiguous anteriorly, diverging posteriorly (length of series, 8.5
mm.; posterior width, 6 mm.) separated from vomerine series by a
distance more than a third the distance between choanae. A very
large hedonic gland on chin, 8.2 mm. wide.

Skin of body smooth dorsally and ventrally, slightly wrinkled
laterally, the pits more or less evident over most of body; costal
folds, 11, not counting an axillary which is not apparent in speci-
mens examined; a strong fold on neck, with lateral grooves which
meet at an angle on neck; vertical groove at angle of mouth which
can be traced across throat; a strong longitudinal fold from eye to
nuchal groove; tail subcircular in outline not, or scarcely, compressed,
the vertical grooves on sides fairly distinct (22 in evidence), the
dorsal surface corrugated and roughened, below perfectly smooth;
the complete but dissevered tail measures 110 mm., which is exactly
the measurement from tip of snout to the back part of vent. (In
younger specimens tail distinctly shorter than body, and distinctly
compressed laterally, with tail strongly constricted at base.) Glandu-
lar spot present behind insertion of femur; limbs heavy, the digits
terminating in the calloused tips which give them a somewhat
truncate appearance; ascending order of size of fingers, 1, 4, 2, 3;
fingers free save for a very minute but distinct web; first finger
smallest and very short l:)ut definitely extending beyond web; order
of size in toes, 1, 5, 2, 4, 3; first toe is very narrow and sliort, only
extreme tip free. Arm brought forward; the longest finger reaches
the posterior corner of eye; when limbs are adpressed the toes are
separated by the width of a costal fold (in large female by about
three folds); lateral fold caused by extension of a hyoid cartilege

268 The University Science Bulletin

terminates at about third or fourth costal fold; sides of anal slit,
with numerous papillae (in females sides of slit folded).

Color in alcohol. Above slaty to grayish-black, of a somewhat
lighter shade below and somewhat darker laterally; a small V-shaped
orange spot behind the groove crossing neck followed by 15 pairs of
orange spots on back which terminate abruptly at tail; latter uni-
form gray slate; tips of digits somewhat lighter than remainder of
hands and feet.

Variation. In MCZ 8436 5 the dorsal spots are less distinct
posteriorly and are smaller throughout with less definite outlines;
there are traces of yellow flecks anterior to the dorsal nuchal groove,
while dorsal and lateral surfaces of the head have numerous brown
spots; a few spots evident on back. A younger specimen, MCZ No.
8437, likewise shows the minute brown spotting. This latter speci-
men has the first three orange spots on each side confluent and the
pairs of dorsal spots are closer together, anteriorly, than in type;
the dorsal ground color has more brown and the ventral surface is
a dull grayish-brown. The pigment is less dense on venter, showing
a cream ground color, especially under hind limbs. The tail, how-
ever, is slate-black above and below in sharp contrast to body color.
A young specimen (MCZ 8434) has the entire dorsal series more or
less confluent, forming two irregularly-edged stripes on the back. In
a very young specimen, EHT-HMS No. 12085 (snout to vent, 9
mm.), the spotting is scarcely visible; the head is rugose, the color
blackish.

Variation in measurements and proportion are shown in the table.

Relationship. The relationship appears to be with 0. bellii, 0.
robustus, and 0. schmidti, the first two being characterized by
orange dorsal spots or flecks. It has obviously been confused with
0. bellii in the literature. The most salient characters by which the
two may be separated is the heavier pitting of the skin, weak or
wanting in 0. bellii, the proportionally wider head in specimens of
equal length (or very much greater body length in specimens of
equal head size) ; large series of teeth (maxillary, mandibular and
vomerine) in adults of giganteus and the absence of the large oc-
cipital yellow or orange spots. It appears to reach a distinctly larger
size than 0. bellii.

It has been impossible to separate completely the literature ref-
erences to 0. bellii, that may refer to this form. I suspect that this
species is confined to the eastern part of the plateau, while 0. bellii
has a much wider distribution on the plateau. Specimens mentioned

Taylor: Mexican Salamanders

269

by Giinther (Biol. Cent. Amor. 1901, p. 299) from Omilteme in the
Sierra Madre del Sur in Guerrero should be carefully reexamined,
since it is possible that still other species are masquerading under
this name.

Table of measurements in mm. and data of Oedipus giganteus

Number.

Museum

Sex or age

Snout to back end of anal slit.

Length of snout

Snout to arm insertion

Head length to jaw angle

Head width

Eye length

Axilla to groin

Arm

Leg.

Tail

Costal grooves

Legs separated by costal folds. . .
Head width in head-body length.
Maxillary-premaxillary teeth. . . .

Vomerine teeth

Mandibular teeth

8436

MCZ

9
128.5

r^

.38.4

22.. 5

21
7

66.5
26-26
26-26

11
3

6.01
67-68
29-30
67-?

8435

MCZ

&
110
4.8
34.2
19
17
4.9
56
24-26
26 . .5-27

11
1

6.4
49-47
25-25
62-62

8431

MCZ

73

3.3
23
14
12.5

4.4
34.2
17
18
54
11

y2

5.8
44-44
22-22
42-43

8434

MCZ

:^'g-
46

3
13.5
10.2

9.2

3.6
22.5
12

12.6
23.5
11

0

5
3.5-39
16-14t
34-35

12085

EHT

>'g.

19
1.5
7

4.5
3.7
1.5
9.5
5.8
6

11.6

11

Overlap

one ff>!d.

* This dissevered tail is in the jar with two tailless specimens. It appears to belong to
No. 8435, but if it should actually belong to the other, No. 8436, it would show that the
tail is not as long as bodv.

t Tooth row very irregular.

Oedipus smithi sp. nov.

(Plate XXV, figs. 5, r,)

Holotype. EHT-HMS No. 3966 $ , collected by Hobart M.
Smith, Cerro de San Luis, 15 mi. N. W. Oaxaca, Oaxaca, Aug. 5,
1935.

Pcn-atypes. 3965-3969 same data as type. Nos. 15616-15641,
Cerro San Felipe, 15 mi. N. W. Oaxaca, Aug. 20, 22, 1938, Taylor.

Diagnosis. A large salamander, dark brown above; below slaty
gray, with a tinge of red on lighter brown at sides ; 13 costal grooves,
those in axilla and groin rather indistinct; the edges of the grooves
darker so that there appear to be 13 dark vertical stripes on side;
3^2 to 4 costal folds between adpressed limbs; vomerine teeth in

270 The University Science Bllletin

two long, greatly curved series of 12-14 teeth, beginning 1.5 mm.
behind a line drawn between posterior edges of choanae and curving
up and out beyond the outer edges of choanae ; parasphenoid teeth
in two series, narrowly separated anteriorly, but separated by 1.8
millimeters posteriorly; each series six millimeters long.

Description of type. Head rather broad, lacking canthus; nostrils
at tip of snout which is truncate; snout extending slightly beyond
mouth (.8 mm.) (in male, 1.5 mm.) ; upper surface of snout slightly
convex; a strong medial groove begins at a point between eyes and
passes back on neck; inner border of eyelid bordered by a distinct
curving groove; parietal regions swollen strongly; head length in
snout to vent length, 5.6 times; head width in snout to vent length,
714 times; eye to tip of snout (3.1mm.) much less than length of
eye (measured from corners of lids, 4 mm.) ; outline of upper jaw,
slightly undulant, seen from side, not straight; angle of jaw much
behind posterior angle of eye, both eyelids fitting under a fold of
skin behind; a groove from eye curves back and somewhat down
along side of neck to beyond the gular fold; first gular groove
rather indistinct, somewhat behind jaw angle; arm well-developed,
the digits webbed for nearly a third of their length, the web con-
tinued to tips as a slight dermal fringe; the first finger very short,
not completely involved in web; the descending order of length of
fingers, 3, 2, 4, 1, the second and fourth of nearly equal length;
toes with a small web at base, and lateral dermal fringe to tips;
3, 4, 2, 5, 1, the order of length of toes; tail (76 mm.) longer than
snout-to-vent measurement (68 mm.), constricted at base, com-
pressed somewhat laterally; anal lips with grooves and folds, with-
out papillae; a well-defined gular fold; vomerine teeth in two curved
series each consisting of 12 (13) teeth, curving forward and out
beyond outer edge of the choanae, separated by 1 mm. from the
parasphenoid teeth; latter in two series beginning at hinder level
of maxillary teeth, narrowly separated anteriorly where they are
very narrow, widely separated posteriorly where the series is widest;
about 19 diagonal rows in each series; 20-22 maxillary teeth; six
or seven premaxillary teeth; 24 mandibular teeth.

Color. Above dark brown; sides somewhat yellowish or grayish
olive-brown tinged with red in life, with a series of thirteen vertical,
blackish bars on side between axilla and groin; chin and under tail
("ream with a peppering of cinnamon-brown; belly grayish-cream
with traces of darker lines following costal grooves; sides of tail
with numerous blackish spots; lips and side of neck cream, peppered
with cinnamon-brown.

Taylor: Mexican Salamanders

271

Variation. There appears to be practically no variation of im-
port as regards color and markings; No. 3965 has the lateral ver-
tical black bars continued across abdomen. In the others the ven-
tral bars are only faintly indicated.

All the males have a very clearly defined gland on the anterior
part of chin which is flat, somewhat salient, 4.5 mm. long, 5 mm.
wide. This is almost without pigment and is in strong contrast to
its surroundings. In the males there are 4 to 6 premaxillary teeth
which pierce the upper lip; the total number of teeth, however, is
about the same as in the female; fourteen is the maximum number
of vomerine teeth. The tails on the males are somewhat thicker,
heavier and a little shorter; the anal walls are strongly papillate;
the snout projects 1.5mm. beyond mouth in the larger males; the
head is proportionally wider in males. The head length, in snout-
to-vent length, being 5; the head width in same being 6.3 times, in
No. 3970. A semicircular fold under tongue of all.

Table of measurements of Oedipus smithi sp. nov.

3966

3965

3970

3968

3969

Sex

9

69
76

69
70

72
69.5

67.5
70.5

cf

59

Tail

66.3

Head to arm .

20
12

22
14.5

22.2
14.8

21.8
15.4

IS. 3

Head length

12.5

Head width

9

11

12

11.2

9

.•\rm

13.5
16

13.6
15.4

14.9
16

13.1
16.5

13.3

Leg

14.3

Axilla to groin

39

38.3

42

38.5

33.3

Interorhital width

2.5

2.2

2.4

2.2

2

3

4

3.8

3.7

2 9

Re^narks. This entire series of specimens was collected by Dr.
Hobart M. Smith. The following data are from his diary. Color
in life: "Below, dull-colored, slaty-gray; above very dark brown,
lighter on sides witli a tinge of red. All were found under planks
and cut logs, covered with leaves, but never very deeply covered.
Forest with many pines and other evergreens, with much moss on
the trees — a sort of rain forest, very damp. Taken about 200 feet
from the summit of the peak of Cerro de San Luis, about 15 miles
(by the road) north of Oaxaca. The elevation must have been
about 9,000 or more feet."

272 The University Science Bulletin

Apparently the species is not closely related to any other Mexican
Oedipus, with the possible exception of 0. sidcatus, from which it
differs in tlie much larger size, longer tail, larger series of vomerine
teeth, a much greater width between adpressed toes and a very dis-
tinctly dift'erent color pattern. 0. sulcatiis has been reported from
Cerro de San Felipe, Oaxaca.

The species is dedicated to Dr. Hobart M. Smith, the discoverer
of this form, in recognition of his researches in Mexican herpetology.

Dunn's reference, of 2 specimens from this locality, to sulcatvs is
not wholly conclusive. However, the comparison given is with his
description, since I have not seen the type. The feet of Dunn's
specimen (U. S. N. M. 47606) differ from Brocchi's figures. (]\Iiss.
Sci. au ^lexiciue. Batrach. Liv. 3.1883 PI. XX, fig. 2.1

Oedipus altamo)dani<s sp. nov.

(Plate XXV. figs. 3, 4)

Holotype. EHT-HMS No. 12245; collected at Lake Zempoala,
Morelos, Mexico, Aug. 6, 1936, by E. H. Taylor; elevation 10,500
feet.

Paratype. EHT-HMS No. 12239. Same data as type.

Diagnosis. A medium-sized species with a rather large head;
vomerine teeth in two series of 10 teeth rather widely separated
medially, curving, extending beyond outer level of choanae; maxil-
lary teeth large, about 22 in maxillary-premaxillary series; mandib-
ular teeth same; hedonic gland on chin of males scarcely discerni-
ble externally; arms and legs well developed, touching when
adpressed; a trace of a web; 12 costal folds, that in axilla very in-
distinct. Purplish with cream flecks on body; head width in snout-
to-vent length, 6.07; head length in same, 4.7; snout to vent, 48.6
mm.; tail, 47 mm.

Description of the type. Head flat between orbits, but surface
somewhat roughened, the snout rounded in dorsal profile; canthus
rostralis lacking or greatly rounded; a slight depression from an-
terior angle of eye toward nostril, below which the lores are more
rounded; nostrils almost terminal, the distance between them about
ecpal to the interorbital width; width of eyelid a little less than
interorbital distance (eciual in paratype) ; posterior ends of eyelids
fitting under a fold; length of eye greater than length of snout; a
deep longitudinal groove from eye back to the ends of the well-
defined gular fold; first nuchal (or head) groove crosses behind
angle of mouth, intersects the longitudinal groove, and is continued

Taylor: Mexican Salamanders 273

some distance above, not crossing chin below; 12 costal folds, that
in axilla dim, none indicated in the groin; a slight indication of a
broken median dorsal groove not reached by the costal grooves; 9
costal grooves continue across belly; anal region much swollen, the
papillae showing deep in the cloaca ; arms and legs strong, elongate ;
very slight webbing indicated; jfirst finger very short, the tip barely
free; fourth finger shorter than second; third much longer than
second; first and fifth toes very short on left foot, on right the fifth
is considerably more elongated; the order of size is 1, 5, 3, 4, 3. A
slight web is indicated; tips of digits swollen on under surface;
skin of head pitted, minutely corrugated; on body, back rather
smooth, the sides wrinkled, belly smooth with fine transverse grooves
or wrinkles; hedonic gland barely visible externally; a small glan-
dular area present behind and a little above insertion of hind leg.

Vomerine teeth in two curved series of 10-11 teeth separated by
a distance equal to one and one half times the width of a choanae,
extending much beyond outer level of choanae; palatine teeth in
two series contiguous anteriorly, diverging and widening posteriorly,
separated from the vomerine series by a distance equal to three or
four times width of choanae ; three or four premaxillary teeth pierce
the lip; 22-23 teeth in maxillary-premaxillary tooth series; about
same number of mandibular teeth.

Color. Head and body generally purplish-violet with a lighter
brownish-lavender clouding along the dorsal surface; cream-laven-
der on sides; ventral surfaces nearly uniform purplish violet (para-
type has creamy flecks on chin and breast, scarcely discernible in
type) ; on the dorsal surface of the proximal parts of the limbs are
creamy areas or spots, these are less distinct on the sides and dorsum
of the distal part of fimbs; lower eyelid grayish.

Measurements {in mm), and data of the type and paratijpe, re-
spectively. Numbers 12245, 12239; sex, ^ , 5 ; snout to vent,
48.6, 40; snout to arm, 15, 14.1; axilla to groin, 27, 23; width of
head, 8, 7; length of head, 10.2, 8.5; foreleg, 13, 11.3; hind leg, 16.6,
12; head width in snout-to-vent length, 6.07, 5.7; head length in
same, 4.7; 4.7; length of eye, 3, 2.6; length of snout, 2.8, 2; in-
terorbital width, 2.2, 1.9; width of eyelid, 2, 1.9; distance between
nostrils, 2.3, 2; snout to, gular fold, 11.6, 8.9; tail, 47, (?).

Remarks. This species belongs in the group having fingers nearly
free, which includes leprosvs and robertsi, but differs from these
species in the larger maxillary teeth; the reduced number in the
maxillary-premaxillary series; the longer, stronger limbs; the re-

18—4141

274 The University Science Bulletin

diiced condition of the fifth toe; and the peculiar coloration clearly
distinguish this species from other members of the genus.

The specimens were collected at about 10,500 feet elevation. I
suspect that it will be found to be a species restricted to the highest
peaks in the region.

The types were found on grass which was partly covered by logs.
In life the violet-purple color with the cream markings and mottling
is very striking.

Oedipus leprosus (Cope)

(Plate XXIX, fig. 2)

1869. Spelerpes leprosus Cope (part.), Proc. Acad. Nat. Sci. Philadelphia, 18G9, p.
105-106 (type description; type locality, Orizava, Mexico, F. Sumichrast, coll.).

1893. Spelerpes gibbicaudiis Blatchley, Proc. U. S. Nat. Miis., XVI, 1893, pp. 38-39
(same type as S. leprosus).

1926. Oedipus cephalicus Dunn (part.), The salamanders of the family Plethodontidae,
Smith Coll. Publ. 1926, pp. 380-384, (USNM 19255; type of gibbicaudiis; MCZ 7659
Xometla, Gadow, coll.).

(I have not attempted to allocate all the various literature ref-
erences to 0. leprosus, since without an examination of the material
on which each report is based such allocation would perforce have to
be made largely on conjecture.)

A reexamination of Cope's type and paratypes of Spelerpes
leprosus have led to the surprising discovery that no less than four
species were present in the original lot. Of the original six speci-
mens, five still remain; the smallest, mentioned in the type descrip-
tion, is lost. ("Like other Mexican Spelerpes, this animal seems
to pass its metamorphoses early; a young one sent with the adults
measures 21 lines [about 46 mm.] in length.")

In the original type description, which is a composite one, Cope
designates as a type a specimen whose measurements are given. This
specimen now bears the number USNM 19255, and is likewise the
type of Spelerpes gibbicaudus Blatchley. This name, therefore, is a
synonym of 0. leprosus (Cope).

Among the four remaining, one (now USNM 123591) is a speci-
men belonging to Oedipus cephalicus (Cope). The identity of an-
other (now USNM 123592) is still in doubt. The two remaining
specimens still bearing the USNM No. 6340 are conspecific and
may belong to an undescribed form.

The exact elevation on Orizaba, at which the type was collected,
is not known. Sumichrast states that it was confined to the "Alpine
region." I have recently examined a specimen which I believe be-
longs to this species (MCZ No. 7659, collected by H. Gadow at

Taylor: Mexican Salamanders 275

Xometla, Camp) which was obtained at an elevation of 10,000
feet on Mt. Orizaba. It presents the following characters:

Adult male. Head not flattened, the dorsal surface of snout curv-
ing, sharply truncate in front, somewhat angular, due to moderately
prominent subnarial swellings; eyes very prominent; body more or
less cylindrical; pitting on skin dim, the skin more or less corru-
gated; the corrugations distinct behind eye; length of eye (3.2mm.)
a little less than length of snout (3.7 mm.); interorbital distance
(2 mm.) about equal to width of upper eyelid (2.1mm.); distance
between nostrils, 3.2 mm.; between choanae, 2mm.; head width
(7 mm.) contained in snout-to-vent length (57 mm.) 8.1 times; head
length (11mm.) in same distance, 5.1 times.

Skin above generally smooth, with very fine, short wrinkles visible
under the lens; sides and venter smooth, the 11 costal folds moder-
ately distinct, the axillary and' inguinal folds not apparent; a whit-
ish glandular area behind insertion of femur ; neck rather constricted
(normal?), the nuchal fold present, with lateral grooves meeting at
an angle on dorsal side of neck; a groove passing across angle of
jaws cannot be traced completely across chin; a well-defined groove
from behind eye to nuchal groove; skin bordering this groove some-
what pustular; anal slit bordered with papillae.

Limbs well developed, but separated when adpressed by a dis-
tance equal to slightly less than width of three costal folds; ascend-
ing order of length in fingers, 1, 4, 2, 3; the digits wide with a
distinct but very short web; tip of first finger free; web involves
about half of the proximal phalanges and is continued along the
edge somewhat on the middle finger; ascending order of size in
toes, 1, 5, 2, 4, 3, the web slightly evident, involving half of the
proximal phalanges on third and fourth toes; tips of digits moder-
ately inflated, but not widened at tip; arm, brought forward, fails to
reach the eye; tail missing. A large hedonic gland on chin 3.2mm.
by 2.5 mm.

Parasphenoid teeth in two groups, separated throughout, diverg-
ing and widening somewhat posteriorly; length, 5 mm.; combined
posterior width, 2.4 mm., separated from vomerine teeth by a dis-
tance equal to half the distance between choanae; 13-14 vomerine
teeth in nearly transverse series, curving back slightly, medially,
separated by a distance not larger than diameter of choanae; a
small pit between choanae; 36-37 maxillary-premaxillaiy teeth;
about four premaxillary teeth pierce lip ; 42 mandibular teeth on one
side.

276 The University Science Bulletin

Color in alcohol. Above, ground color slate with grayish cloud-
ing or blotching on dorsal surfaces and sides; laterally the costal
grooves show as brownish lines; uniform slate below; limbs lighter
with grayish blotches; underside of hands and feet yellowish-cream;
snout creamy with thin scattering of pigment, absent or nearly so
on subnarial swelling and lips; lower eyelid cream edged with
black; upper eyelid edged with cream; chin largely cream with
minute scattering of darker pigment and a cream spot across throat
at fold.

Measurements. Snout to posterior border of vent, 57 mm. ; snout
to arm insertion, 17 mm.; axilla to groin, 30.4 mm.; head length,
11 mm.; head width, 7 mm.; arm, 11.8 mm.; hind leg, 13 mm.

Variation. The description of Spelerpes gibbicaudus by Blatchley
offers some details on the type of Spelerpes leprosus Cope. My
examination of the type shows the proximal phalanges of the middle
toes and fingers to be involved nearly half their length in the skin
(web) ; the groove behind the eye is only dimly visible. The type
description by Blatchley states that the toes are unwebbed. The
pitting on the skin is minute and minute wrinkles are visible.
Twelve costal folds can be counted; 24 grooves visible on tail;
maxillary-premaxillary teeth, 32-32; vomerine teeth, 12-12. Brown-
ish coloration mentioned by Blatchley is apparently due to pres-
ervation. Chin cream with a meager scattering of pigment; head
lighter than body, the tip of snout nearly all cream; lips cream
and eyelids with some cream color; below, brownish slate; grayish
on top of tail and dark below.

Measurements (in mm.) of the type of Spelerpes leprosus given
by Cope (reduced to millimeters) ; of the same specimen as given
by Blatchley in his description of Spelerpes gibbicaudus; and my
measurements of the same specimen, respectively: Snout to pos-
terior end of vent, 50.8, 46 (may be only to front end of vent),
48.5; total length, 88.9, 85, 87; tail, 38.1, 38.5, 39; axilla to groin,
29.4, 26.3, 31; arm, 10.5, 10,- 10; leg, 10.9, 10.7, 11; head width, 7.6,
7, 6.33 (this measurement varies if the mouth, which has been
forced open, is not completely closed).

I suspect that the slight differences in measurements are due
merely to different techniques in measuring. The specimen now has
the tail severed. None of the other specimens in the series ap-
proaches closely the measurements given by Cope for the type.*

♦Certain workers have stated that they do not believe a type has been indicated by
Cope. While I regard the act needless, I hereby designate the presumed type (now, USNM
No. 19255) as lectotype.

Taylor: Mexican Salamanders 277

Oedipiis manni sp. nov.

(Plate XXIX, figs. 4, 5, 6)

1918. Oedipus leprosus Dunn, Bull. Mus. Comp. Zool., G2, 1918, p. 470. (Harvard
specimens 3912-3930), Guerrero, Hidalgo, Mexico.

1926. Oedipus cephalicus (part.) Dunn, the salamanders of the family Plethodontidae.
Smith College publ. 1926, pp. 380-384 (MCZ Nos. 3912-3929).

Type. MCZ No. 3915, Guerrero, Hidalgo, Mexico; W. M. Mann,
collector.

Paratypes. MCZ Nos. 3912-3914; 3916-3927. Michigan U. mu-
seum, Nos. 48061-48062, type locality, W. M. Mann, collector;
EHT-HMS, Nos. 15656-15657, near Zacualtipan, Hidalgo.

Diagnosis. Belonging in the cephalicus group, with well-developed
limbs and partially webbed feet. The first finger and toe slightly
emergent from the "web"; outer toe extending distinctly beyond
the ''web"; limbs, when adpressed, separated by about two costal
folds; dorsal surface of head with very fine pits; back smooth, the
pits obsolete; about 38-44 teeth in the maxillary-premaxillary series
of females; 33-35 males; 13-16 vomerine teeth, each series extending
beyond nares and curving back to meet at a point medially one
millimeter behind their anterior edges; snout moderate in females,
sharply truncate with slight emargination, and two greatly-developed
subnarial knoblike tubercles in males; head width in snout-to-vent
length, 5I/2 to 61/0 times; a groove from eye to gular fold; 12- (13)
costal grooves. Black or brownish-black with silver spots or flecks,
more prominent on sides and belly; throat black with silver or
cream flecks.

Description of the type. Head moderately flattened; eye
(3.05 mm.) much longer than length of snout (2.1 mm.), slightly
longer than its distance from middle of the tip of snout. Width of
an upper eyelid equal to the interorbital distance ; distance between
nostrils, 2.15 mm.; eyelids with their posterior parts fitting under
a diagonal skin fold; very small pits indicated on the top of head,
distinctly larger and closer together on eyelids and temporal region ;
maxillary-premaxillary tooth series, 44-44; 13-14 vomerine teeth,
extending beyond the very minute choanae (.018 mm.), curving
back and terminating slightly more than a millimeter back of their
anterior border; the two vomerine series separated by a distance
about equal to the diameter of the choanae; palatine teeth in two
club-shaped series scarcely separated anteriorly, but distinctly
diverging posteriorly, the total length, 4 mm.; width, 2.4 mm. pos-
teriorly; separated from the vomerine series by a distance of one
millimeter; mandibular teeth, about 40-40.

278

The University Science Bulletin

Body apparently somewhat compressed and slightly elevated
along middle part of body ; 12 costal folds not counting an inguinal
fold, which apparently is obsolete or absent; skin on sides wrinkled
slightly or minutely corrugated; on back smooth, on belly smooth
with fine transverse lines, two or three to each fold; tail rather
strongly constricted at base.

The vertical tail grooves are not strongly marked except on
proximal portion; tail more or less wrinkled everywhere, somewhat
compressed laterally ; length of tail about one fifth less than head-
body length; limbs moderately heavy, the first toe short, only the
extreme tip extending beyond level of web ; between the two middle
fingers the web includes more than half of the proximal phalanges;
fourth finger about one millimeter long; the web appears to continue
along edge of digits as a fringe if the hand is slightly dried; fingers,
3, 2, 4, 1 in descending order of size; arm brought forward, the long-
est finger reaching posterior corner of eye; foot with a distinct web,
including half the proximal phalanx of the second and third toes,
all of the proximal phalanx of the fourth and fifth toes; first very
slightly emergent from web; fifth toe extends beyond web one half
millimeter or more; toes, 3, 4, 2, 5, 1 in descending order of size: ad-

Measurement.-? in mm. and data from Oedipus manni sp. nov.

Number

3915

3920

3925

3916

3923

3912

Museum

MCZ

MCZ

MCZ

MCZ

MCZ

MCZ

Sex

9

9

9

(ji

&

?

Snout to back end of anal slit

53.2
2.1

52

46
2 2

48
3

43

2.7

39 5

Length of snout

1.9

Head length to jaw angle

10

8.2

8.8

8.2

7.2

Head width

8.4

8

7.8

7 4

6 8

Eye

3.05

2.9

2.5

3

2.5

Axilla to groin

28

30

23.5

22

2''

22

Arm

12.4

11.2-12

11

13

12

9 8

Leg

13.2

12.5-13

11.2

12.3

n .5

Tail

41

36

32

40 2

34

27 5

Costal grooves

12

13

12

13

12

12

Legs separated by folds

2

2y2

2

touch

touch

2

Head width in head-body length

6.1

5.75

6.15

5.81

56.2

Maxillary-premaxillary teeth

44-44

40- ?

38

3.3-35

35-35

34-34

Vomerine teeth

13-14

13-13

13-13

?

15-16

15-14

Mandibiilar teeth

40-40

39-40

40-41

42-41

36-35

36-38

Taylor: Mexican Salamanders 279

pressed limbs fail to meet by a distance equal to width of two folds ;
anal slit, 4mm. long, the sides with diagonal folds; a strong gular
fold across neck from which ascends a groove which cannot be traced
to the dorsal surface; a groove crosses chin and reaches up beyond
angle of mouth; a small glandular spot present behind insertion of
femur.

Color (in alcohol). Dorsal surface grayish-brown (reddish-brown
in life?), the head slightly darker; sides growing slightly darker
towards venter; sides with a frosting of silver growing slightly more
dense low on side; it is then replaced by scattered flecks and spots
of cream or silver on ventral part of body and head; the chin region
has the ground color darker and the silver spotting more dense
than on abdomen; tail and limbs dark, flecked or spotted with
silver; lower eyelid with some cream or yellow color.

Vanatioji. In the paratype series, Nos. 3913, 3917, 3918, 3919,
3921, 3923, 3924, 3926, have been preserved in a different manner
from the remainder of the MCZ specimens of the series. The ground
color appears to be a deeper black and the frosting of the dorsal
surface is obscured. The contrast of the black and silver on the
belly is very pronounced. The specimens are somewhat shrivelled
(as are specimens placed in too strong a solution of alcohol). The
remainder of the series is well-preserved and the general appearance
of the specimens (under water) is grayish on the sides and a dull
black, grayish-black or grayish-brown along the back; the front
borders of the eyelids and the tip of the snout are lighter, and the
loreal region slightly darker than remainder of head; lips heavily
flecked with whitish silver. The ventral ground color is a dull
grayish black with the- silver spots and flecks in strong contrast.
Tail with silver flecks generally scattered or segregated, leaving
large blackish spots (No. 3912) on tail.

The principal differences in the specimens are sexual, the males
having a longer, very truncate snout, slightly emarginate when
viewed in profile from above; the under side of the snout, which
projects strongly beyond the mouth, is concave between the two
greatly developed subnarial, knoblike bosses which project at the
angles of the snout; the legs are a trifle longer and touch (at least
in adult specimens) when adpressed.

A well-developed hedonic gland is present on chin of males. The
eye is longer than the snout in females, nearly the same length or
shorter in males; anal lips with papilla in male, and the premaxil-
lary teeth (not more than four) pierce the lip. The reduction in

280 The University Science Bulletin

the number of teeth in the adult male is another sexual character.
Older females seem to have a larger series of teeth than younger
specimens.

Rernarks. A male and female of this form (the latter the type)
are described as Oedipus cephalicus by Dunn {loc. cit. 1926, pp.
381-382). The relationship of the species is with cephalicus. It
differs strikingly in coloration and marking, the color of cephalicus
being a dull blackish to bluish or grayish slate without distinct
markings on body and tail; the tail of 0. cephalicus is slenderer,
less compressed, tapering more gradually and is distinctly longer;
the axilla to groin measurement is longer and the limbs likewise
longer than in specimens of 0. manni of the same snout-to-vent
length. The subnarial bosses are proportionally larger and the
snout narrower and the pitting on head and body less conspicuous
in 0. manni; and the body is slightly compressed and deeper in the
middle than in 0. cephalicus.

The species is named for Dr. W. M. Mann, director of the Na-
tional Zoological Gardens in Washington, the discoverer of the
species. I am indebted to Dr. T. Barbour and Mr. A. Loveridge for
the privilege of describing the species.

Oedipus orizabensis (Blatchley)

(PI. XXV, figs. 1, 2)

1910. Spelerpes orizabensis Blatchley, Proc. U. S. Nat. Mus., 1893, p. 37 (type de-
scription; type locality, Mt. Orizaba, Viracriiz, Mexico); Gadow, Proc. Zool. Soc. London,
1905, p. 203; Zool. Jahrb., 1910, pp. 709, 714.

1926. Oedipus cephalicus Dunn, The salamanders of the family Plethodontidae, Smith
College Pub., 1926, pp. 380-384 (part.).

The species is represented in the collection by the following speci-
mens EHT-HMS, Nos. 4000-4011, 4013-4023, 4025-4116, 4118-4123,
4125-4126, 4128, 4384 (Taylor-Smith), July 30-31, 1932, and 12188-
12212, 12214-12216, 12281-12223, 12227, Aug. 12, 1936 (Taylor),
between kilometers 58-66, west of Rio Frio, Puebla, elevation, 8,000
to 10,000 feet. No. 12225, near Las Vigas, Veracruz, elevation about
8,000 feet. Sept. 1, 1936 (Taylor). Nos. 4327-4367, 4378-4379, July
19, 1932 (Taylor and Smith), and 12044, 12045, 12048-12051,
12054, 12056-12058, 12062-12064, 12066, 12068, 12070, 12075, 12078-
12080, 12083-12084, 12086, 12088, Sept. 2, 1936 (Taylor), Cruz
Blanco and slopes of Cofre de Perote up to about 11,000 feet. Nos.
12099-12108, 12110-12113, 12115-12118, 12121-12122, 12232, 12235,
12236, 12241, 12244, 12247-12252, 12255-12257, 12259, 12263, 12265,
12266, Lake Zempoala, Morelos, 10,000 feet, Aug. 6,7, 1936 (Taylor).
Nos. 12267-12268, 12412, 12415, 12416, in a pedrigal, at kilometer 35

Taylor: Mexican Salamanders 281

on road between Mexico City and Trcs Cumbres, Morelos, July 8,
9, 1936 (Taylor and Smith).

Diagnosis. A grayish-black, medium-sized species with a tend-
ency to grayish clouding on back; chin lighter; head flat; no
canthus; palatine teeth in two slightly diverging series; separated
from the vomerine teeth (rarely continuous) ; vomerine teeth, 12-17
in a curved series extending beyond choanae; adpressed limbs sepa-
rated by from 1% to 4 costal folds; costal folds, 12-13; tail as long
as head and body in adults; a small gland (appearing as a whitish
spot) behind and slightly above the hind limb; head width in snout-
to-vent length (above 50 mm.), about 7.5; head length in same,
about 6 times. Hedonic gland on chin in males ; the subnarial swell-
ing much smaller than in 0. cephalicus males, or 0. leprosus males.

Description of the species. (EHT-HMS No. 12049 $ , slopes of
Cofre de Perote near Cruz Blanca, Veracruz; Taylor-Smith, col-
lectors.) Head rather flat in occipital and interorbital regions,
rounding on snout ; no canthus present ; snout bluntly oval, the swell-
ings below nostrils scarcely discernible; length of eye (2.5 mm.)
greater than length of snout (2.1 mm.) ; upper eyelid (1.8 mm.)
less than interorbital distance (2.3 mm.) ; distance between nostrils,
2 mm.; head width in snout-to-vent length, 7.2 times; head length
in same, 6.4 times.

Thirteen costal folds, those in axilla and groin distinct; a discon-
tinuous groove along middle of back not reached by the costal
grooves; first gular groove crosses angle of jaw from about upper
level of eye, and just fails to meet its fellow in middle of throat be-
low; nuchal fold w4th irregular grooves arising from its ends which
reach the median line forming an angle, pointing forward; a deep
groove behind eye intersects the first gular groove and passes back,
becoming continuous with the gular fold; costal grooves continued
across belly ; caudal grooves very distinct in adults ; adpressed limbs
fail to meet by a distance equal to about four costal folds; digits
slightly webbed, the first very short with a free tip; the second is
a trifle longer than fourth, the third longest; finger margined with a
discernible fold of skin, and the ventral surface of the tip is padded;
toes in the following ascending order of size: 1, 5, 2, 4, 3.

End of part of the hyoid apparatus causes a prominent, elongate
raised area from gular fold, passing along the shoulder to a point
some distance behind the arm; a glandular area behind and some-
what above the insertion of leg; skin of head thickly and regularly
pitted as is the skin of dorsal and lateral parts of body; ventral
surface with very minute pits.

282

The University Science Bulletin

Vomerine teeth in two series of 14-15, narrowly separated medi-
ally, curving, extending far beyond outer level of the choanae, sepa-
rated from the palatine teeth by a distance equal to four or five
times the diameter of the very small choanae; palatine series dis-
tinctly separated, diverging posteriorly the space between their
posterior ends nearly double the width of one series; about forty
maxillary teeth, those on premaxilla alternately large and small;
41-42 mandibular teeth.

Colo7\ Generally grayish-black to slate, flecked or clouded above
on body and tail with grayish-brown, and on sides with grayish-
cream; below plumbeous, the throat and chin yellowish-white with
scattered peppering of black; lower eyelid whitish; a cream mark
on the tip of snout resembling an inverted U or V; gland behind
and above insertion of hind leg grayish; underside of hands and
feet dirty whitish; tiny arrow-shaped cream spots near tlie tips of
the digits.

Table of measurements (in mm.) and data on Oedipus orizabcnsis Blatchley

Number.

Sex

Snout to vent

Snout to foreleg

Axilla to groin

Head width

Head length

Tail

Arm

Leg

Vomerine teeth

Costal grooves

Head width in Igth. (times),
Head length in Igth. (times).

4000

12048

12049

12268

4379

12069

9

9

9

9

&

cf

55

57.5

51

56

53

52

14.9

15.8

14

15

15

16

35.6

34.5

30

33.8

32.2

28

7.3

7.2

7

7.2

7

7

9.1

8.3

7.9

9.2

8.7

8.9

45

51

51

55

53.3

54

10.5

11.4

8.5

11.7

11.5

10.8

10.5

12

10

12

13.4

12.3

17-17

13-14

14-15

1,5-17

12-13

13-13

13

13

13

13

13

13

7.5

8

7.28

7.7

7.5

7.4

6.04

6.9

6.4

6.1

6.1

5.9

4024

&
45.4
14
27.5

6.5

7.8
51
11
11

13-13
13

7

5.8

Variation. Some variation is observable in the number of vo-
merine teeth and the relation of the series to the palatine tectli. The
maximum number of vomerine teeth observed was 17, and in this
specimen, EHT-HMS 4000, the palatine and vomerine series were
practically continuous. One other specimen had one or two teeth
between the median ends of the vomerine series, making them
practically continuous. The palatine series are about 5 millimeters

Taylor: Mexican Salamanders 283

long. In EHT-HMS 4067 the scries are together anteriorly and
diverge posteriorly the width of one series.

Males have a well-developed hedonic gland on the tip of the chin;
the premaxillary teeth (usually 2) pierce the lip, and occasionally
are visible when the mouth is normally closed.

Specimens from Cruz Blanca and the slopes of Cofre de Perote
often have brownish fiecks and clouding on the dorsal surface of
back and tail and the digits are a trifle larger, the hand and foot
having a little more width; the light figure on the snout is distinct
and in young the tip may be largely cream. The sides are often
"frosted" with silver. Specimens from the region near Rio Frio,
Puebla, have the dorsal surface of the tail with heavier clouding of
cream and the snout marking is discernible only in younger speci-
mens. Those from the Ajusco mountains (region of Trcs Cumbres
and Lake Zempoala) are more plumbeous, the light markings on the
tail not so well pronounced; the dorsal and lateral silvery clouding
is rather equally distributed.

The head width and length, in the snout-to-vent length, varies
with the age and length. The head is contained a fewer number of
times in younger specimens. The number of folds between the
adpressed limbs is likewise variable with age. The axilla-to-groin
measurement is greater in females containing eggs. Tails in younger
specimens vary, being shorter than head and body. There is much
variation in the thickness of the tails of preserved specimens. This
is due to the amount of secretion from the tail.

Remarks. This species may be readily distinguished from Oedi-
pus cephalicus by the larger feet and hands of that species, with
their greater amount of webbing. Differentiation from 0. alta-
montanus and 0. leprosus is discussed under those species.

This seems to be one of the most common species in Mexico. The
number of individuals taken is exceeded only by the smaller 0.
chiropterus. They are found together on the forest floor, under logs
and debris.

I refer the following MCZ specimens to this species: Nos. 8404,
8417, 8418, 8420, 8421, 8423, 8426, 8427, 8431, all from Popocatepetl
volcano, Mexico.

Oedipus cephalicus (Cope)

(Plate XXVI, figs. 3, 4)

186.5. Spelerpes cephalicus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1865, p. 196 (type
description; type locality, "Mexican Tableland," Dr. C. Sartorius, coll.); and 1869, p. 106
(listed only "N. E. Mexico") Suniichrast, La Naturaleza, 1882, p. 79 (listed); Cope, Bull.
U. S. Nat. Mus., No. 32, 1887, p. 8 (listed only); Boulenger, Cat. Batr. Grad. s. Caud.
British Mus., 2d Ed. 1882, pp. 67-68 (redescription from Cope); Brocchi, Mission Scientifique

284 The University Science Bulletin

au Mexique et dans TAmerique Centrale; Etude sur Batraciens, Livr. 3, 1883, p. 109 (by
description from Cope); Cope, Bull. U. S. Nat. Mus., No. 34, 1889, p. 162 (key); Giinther,
Biologia •Centrali-Americana, Rept. Batr., Dec, 1901, pp. 298, 229 (description after Cope);
De Leon, Indice de las Batracios que se encuentra in la Republica Mexicana, 1904, p. 38
(list); Cope, Amer. Nat., Dec., 1896, p. 1022 (distribution).

1924. Oedipw cephalicus Dunn, Field Mus. Nat. Hist., XII, 1924, pp. 99-100 (key)
(part.) and Salamanders of the family Plethodontidae, Smith Coll. Publ., 1926, pp. 380-
384 (.part.); and Proc. Acad. Nat. Sci. Philadelphia, 88, Oct. 20, 1936, p. 471 (Pablillo,
Nuevo Leon) ; ? Wolterstorff, Abh. Ber. Mus. Nat. Heimatk. Natur. Ver, Magdeburg, Bd.
VI, Heft. 2, 1930, p. 146.

Cope has said of this species: "The foiTn of the present species is
more that of Ambystoma opacum, and is the shortest and stoutest
seen in the genus." The type has been lost and most of the speci-
mens listed by Dunn (1926) as 0. cephalicus apparently belong in
other species.

The species is represented in our collection by a series of more
than fifty specimens, with a range including Hidalgo, Central
Veracruz, Morelos, and Puebla, as follows:

EHT-HMS. Nos. 4117, 4313-4326, Cruz Blanca, Veracruz north side of Cofre
de Perote, 8,000 feet, July 13, 1932, Taylor and Smith; 4368-4370 near Tres
Cumbres, Morelos (km. 35 on highway) July 11, 1932, Taylor and Smith;
4371-4375 Cruz Blanca, Veracruz, July 18, 1932, Taylor and Smith; 4534-4540,
between Rio Frio and Puebla, Puebla, July, 1932, Taylor and Smith; 4012,
Rio Frio, Puebla, July, 1932, Taylor; 12092-12095, 12098, 12097, 12260, 12493,
Lake Zempoala, near Tres Cumbres, Morelos, 8,500 to 10,000 feet, Sept. 4-6,
1936, Taylor; 12269-12270, 12413, km. 58, near Tres Cumbres, Morelos, July 10,
1936, Smith and Taylor; 12495, near Minas Viejas, 7,000 feet, near Jacala,
Hidalgo, on highway, July 5, 1936, Taylor; 12042, 12052, 12059, 12065, 12067,
12073, 12077, 12081, 12123. 12089. 12224, above Cruz Blanca on north side of
Cofre de Perote, Veracruz, in pines, elevation 8,000-10,500 feet, Sept. 2, 1936,
Taylor.

Cope's description contains the following data: "Muzzle rounded,
truncate, with obtuse angles at the nares, its length from line con-
necting anterior canthus oculorum equal length of eye. Distance
between these canthus equal from hinder canthus to nares. Breadth
behind orbits equal length of tibia and foot. Muzzle to axilla equals
% distance from axilla to groin. Costal folds {i. e., dorsal and lum-
bar vertebrae) , eleven. Tail swollen, little compressed, constricted
at base. Posterior limb stout, extending to sixth fold from behind;
toes flat, depressed, margined, inner very rudimental. Inner and
outer digits of anterior limb similar; the longest extend to near the
middle of orbit. Series of vomerine teeth nearly straight, not in
contact; a postgular fold. Skin everywhere finely wrinkled. Color
dull black, paler on the sides, lips and gular region minutely mar-
bled with ashen. "Length of rictus oris, 2.75 lines [6.3 mm.] ;

* I am designating EHT-HMS 4372 as a neotype, since (fide Dunn 1926) the type is lost.
The designated specimen is practically the same size as the type. The measurements cor-
responding to the above are, respectively, 6.2 nmi. ; 15 mm.; 37 mm.; 33.8 mm.; 11.3 mm.

Taylor: Mexican Salamanders 285

length to axilla, 6.8 lines [15.6 mm.] ; length to groin, 16 lines [36.8
mm.] ; length of tail, 15 lines [34.5 mm.] ; length of hind limb, 5.2
lines [11.9 mm.].

Description of species. (From EHT-HMS No. 12098 $ , col-
lected Lake Zempoala, Morelos, Aug. 5, 1936, E. H. Taylor, collector.)
Head flat, the canthus well defined in front of eye, but becoming
rounded and disappearing near nostril; snout sharply truncate, ex-
tending a very short distance beyond mouth; tubercular swellings
below nostril near lip; length of eye slightly longer than snout;
interorbital distance one and one half times the width of eyelid;
distance between nostrils about equal to greatest width of an eyelid ;
posterior edges of upper and lower eyelids pass under a diagonal
fold; posterior line of mouth turns up at rictus; first nuchal groove
begins on sides of head, passes down and completely across throat;
a strong nuchal fold, the grooves from the two sides continuing up
on dorsal surface, where they meet at an angle, directed forward; 12
costal grooves, the axillary and inguinal rather dim; skin between
folds forming numerous longitudinal wrinkles (tail somewhat shriv-
elled due to excessive secretion of mucous; in life tail rather plump) ;
tail distinctly constricted at base; the vertical grooves very indis-
tinct on tail; costal grooves discernible across belly; skin on dorsal
and lateral surfaces and on breast strongly pitted.

Limbs well developed, the anterior brought forward, the finger
reaches to near middle of eye; adpressed limbs separated by less
than two folds (about ly^ to 1% folds) ; fingers and toes partially
webbed, this webbing much thickened, the digits themselves flat-
tened and more or less definitely margined; first finger much smaller
and shorter than fourth; ascending order of length of fingers, 1,
4, 2, 3; of toes, 1, 5, 2, 4, 3; walls of cloaca heavily folded at anal
opening; head width in length (snout-to-vent), 5.9 times; length
of head, in same, 5.3 times.

Vomerine teeth in two nearly straight series of 15-18, which be-
come slightly curved medially, not meeting on median line, the
teeth somewhat irregular; palatine series about 5 mm. long (ab-
normally short on right side) , the groups not tending to diverge pos-
teriorly; separated from the vomerine teeth by a distance equal to
four times the diameter of the very small choanae; vomerine teeth
extend much beyond the outer edges of choanae; maxillary pre-
maxillary series, 39 on each side; 42-45 mandibular teeth.

Color. Blackish or grayish-black above and below, with some
lighter flecks on chin, the flecks less noticeable on abdomen ; tlie un-

286

The University Science Bulletin

derside of tail flecked and clouded with lighter color; upper side of
tail with occasional, very indistinct brownish flecks.

Measurements of Oedipus cephalicus Cope

Number.

Sex

Snout to vent. . . .

Tail

Snout to arm

Axilla to groin . . .
Width of head ...
Length of head . . .

Eye

Eyelid

Snout

Interorbital width .

Arm

Leg

12098

9

53

.50

16.5

.3.3 . 1
9

10
3

2.7
2.9
3.1

13.1

16

4368

52.2

52

16

29.5
8.5

10
3

2.7
3.5
3.1

15.5

16.2

12242

?

57

50

16.5

35.2
9

10
3.5
2.7
2.8
3

15

16.1

4369

60.2

55.5

18

35.2
9.2

11
3.4
2.3
3.2
3.2

15.3

18

4372*

9
40

33.8
13.5
23
7.4
7.1
2.1
2

2.2

2.2

10.9

11.5

* 4372 is designated the neotype.

Variation. The table shows variation in measurement. The tail
is proportionally shorter in younger specimens. Males differ in hav-
ing 33-34 maxillary teeth, with three or four premaxillary teeth
piercing the lip ; 39-42 mandibular teeth ; snout longer than eye ; the
swellings below nostril are greatly inflated, and the snout projects
more beyond mouth; adpressed limbs touch or are more narrowly
separated than in females; males have a flat gland near tip of chin,
and the webbing of the toes is somewhat greater. The smallest
specimen, 18 millimeters, snout-to-vent, has the head greatly rough-
ened, as do others up to 25 millimeters length.

Remarks. The specimens were found under logs, usually not
directly on the earth but on grass or other trash. When captured,
they exuded much mucous secretion, and again this was done when
they were placed in alcohol for killing. The result is that the tails
of all the specimens look shrivelled. The silvery flecks on the ven-
tral surface are occasionally distinct in life, giving the belly a frosty
appearance; this disappears in preserved specimens.

From the various localities there were observable some slight
differences in the webbing of the digits and the spread of hand and
foot. When males and females of equal length are compared the
male usually has slightly larger feet and hands.

Taylor: Mexican Salamanders 287

I have examined certain salamanders in the United States Na-
tional Museum and specimens from the Museum of Comparative
Zoology, Harvard College. I am referring the following specimens to
this species: MCZ Nos. 8408, 8419, 8424, 8428-8430. 9,000 feet
Popocatepetl, Pue., MCZ 8376, Jalapa.

Oedipus rohertsi sp. nov.

(Plate XXVI, fig. 2)

Holotijpe. EHT-HMS, No. 12503, collected Nevada de Toluca,
elevation between 10,000 and 11,000 feet, Sept. 7, 1936; H. Rad-
clvffe Roberts, collector.

Paratijpes. EHT-HMS, Nos. 12496-12498, 12504-12505, collected
same date and locality by Philip Powers, Edwin R. Helwig, Rad-
clyffe Roberts and Edward H. Taylor; Nos. 15600-15615, topotypes,
Taylor.

Diagnosis. A medium-sized species related to 0. orizabensis and
0. leprosus, but differing in having a somewhat more robust body,
with a broad, orange stripe on the back and tail, much larger limbs
and toes, the latter lacking any trace of a web; adpressed limbs in
contact or separated by a part of one costal fold; tail shorter than
or almost equal to head and body length; vomerine teeth, 8 or 9
in a curved series; 13 costal grooves (counting one in axilla which
is very indistinct) ; head length in head body lenglli, 5.7 times; head
width in same, 6.2 times.

Description of type. Head broad, rather flattened; no trace of
canthus; snout truncate, the nostrils very close to anterior point;
snout extending beyond mouth .5 to .7 millimeters; a rounded
swelling below nostril near lip; interorbital width equals distance
between nostrils, a little greater than width of an eyelid; length
of eye about one fifth longer than snout; upper surface of head,
between eyes and on the region behind eyes, flat; occipital region
not, or but slightly, swollen; a strong nuchal fold across the ventral
surface of the neck; a groove from this to dorsal surface, where it
runs forward somewhat, joining the groove from the opposite side;
a short, vertical groove on side of head crosses angle of jaw some-
what back of the angle of the mouth and continues on the side of
head to dorsal surface; a groove from behind eye crosses this groove
and continues back to the nuchal groove, where it terminates; 13
costal grooves, that in axilla very dim; area between grooves very
wrinkled, the upper edge of the wrinkled area suggestive of dis-
continuous sinuous, longitudinal groove; tail constricted at base,

288

The University Science Bvlletin

the folds between the caudal grooves wrinkled; tail compressed
laterally; length of head in snout-to-vent length, 6.2 times; outline
of edge of upper jaw almost straight; posterior ends of both eyelids
fitting under a fold of skin.

Limbs well developed, when adpressed, the digits separated by a
distance of one costal fold or less ; no trace of webs ; digits flattened,
save at tip, which is definitely inflated and rounded; first finger very
short, its tip free; ascending order of length of fingers, 1, 4, 2, 3; of
toes, 1, 5, 2, 4, 3; skin on dorsal surface of head and body more or
less minutely pitted ; tail a little shorter than head and body.

Vomerine teeth in two slightly curved series of 8 or 9 teeth, extend-
ing beyond the outer edges of choanae, separated from each other
by a distance one and one half times the width of a choana ; palatine
teeth in two series, very narrowly separated anteriorly, and diverg-
ing somewhat posteriorly; 19-20 maxillary teeth; 4 or 6 premaxillary
teeth.

Color. Above a broad, variegated, orange-reddish stripe from
head to tip of tail; sides somewhat brownish lavender; below lead
color (specimen is now somewhat discolored and is quite deep brown
on sides and abdomen), a few spots of orange-brown on head and
along sides. Underside of hands and feet immaculate.

Measurements of Oedipus robertsi sp. no\'.

Number

12503

12497

12504

12.505

12498

12496

Sex

9
51

8.2

8.9
17.2
30

2.9

2.3

3

2.3
44.8

&
48.8

9.4

7.8
15
29.2

2.6

2

3

2.4
48.5

9
47.4

9.2

7*
15
26.3

2.9

2.1

2.5

2.6 .
47

9
36.5

7.5

6

11.4
22.2

2

1.8

2.5

2
33

9

38. 1
7.6
6.5

12.9

22.6
2.2
1.5
2.5
2

33.5

9

Snout to vent

34.5

Head length

Head width

7.2
6

Snout to arm

10.5

Axilla to groin

Interorbital width

Eyelid

20.3
2

1.5

2.1

1.9

Tail

30

* Injured.

Variation. In most of the characteristics the paratypes agree with
the type. In the younger specimens the digits touch when adpressed
and the tail may equal the length of head and body. The dorsal
stripe may break up into spots on the tail. A male (No. 12497)

Taylor: jNIexican Salamanders 289

differs in li;i\'ing a lar<j;er swelling below the ndstvils, Hie j^noiit
slightly more truncate, and a well-defined gland on chin near tij)
of the lower jaw; the premaxillary teeth (4) pierce the upper lip and
the anal slit has the wall papillate instead of folded as in the female.
The stripe may be brown or faun.

Rema7-ks. The specimens were found at an elevation Ix'tween
10.000 and 11.000 feet, along the road leading to the summit of the
volcano, Nevada de Toluca. They were found for the most part
under stones, in the ]une forest.

The species is dedicated to INIr. Radclyffe Roberts, of the Phila-
delphia Academy of Sciences, who assisted in collecting the types
and made ]ios<ible the journey to the mountain.

Oedipus multidentata sp. nov.

Oedipuit cliiniptcnis Dunn. Aciui. Nat. .Sfi. Pliihulelphia. 88. 1936, p. 471.

Type. MCZ, No. 14812, ^ ; Alvarez (km. .l^ on Potosi y Rio
Verde R. R.), San Luis Potosi, Mexico, elevation 8,000 feet, W. W.
Brown, collector.

Paratypes. MCZ, Nos. 14810-14811. Topotypes. Brown, col-
lector. UMMZ, Nos. 63946, 63948, 63953; EHT-H^IS, 15658-15848,
El Chico, Hidalgo.

Diagnosi.^. A small species related to Oedipus chiropterus, but
differing in lia\-ing longer and larger limbs and feet which touch or
overlap when adpressed, tail somewhat more attenuated; eye some-
what larger, and head slightly more flattened; adult males with the
maxillary-premaxillary tootli series 20-24 in each half of jaw (in
0. ehJropterus usually 6-6), and tl;e teeth appear to be stouter than
the teeth in the females of chiropterus. Tlie vomerine series assume
a more transverse position, and th.e choanae are somewhat larger.

Description of the type. Head rather flattened; eye 1 2.5 mm.)
slightly shorter than snout (2.7 mm.); distance between nostrils,
2.15 mm.; smallest interorbital width (2.1 mm.) is greater than
width of an upper eyelid (1.7 mm.) ; width of head (6.2 mm.) con-
tained in distance between snout and posterior end of vent (39.5
]nm.) 6.37 times; head length (8.5 nnn.) in same distance. 4.64
times; line of mouth diagonal and somewhat undulant posteriorly;
subnarial swellings proijiinent, giving the snout a slightly angulate
appearance; snout truncate with a ftiintly indicated canthus, below
which is a ^•ery slight depression extending from eye toward nostril;
hedonic gland on chin indistinct. Vomerine teeth, 7-8, extending
near to outer edge of choanae, nearly transverse, but forming an

19—4141

290 The University Science Bulletin

angle posteriorly, the series separated by a distance a little less than
diameter of a choana; parasphenoid teeth in two distinct series,
widening posteriorly and diverging slightly posteriorly, separated
from the vomerine series by a distance equal to two thirds the dis-
tance between choanae; maxillary teeth extending back past the
middle of the eye; premaxillary teeth somewhat more elongate and
slenderer, none piercing the lip but extending outside the lower lip,
the combined maxillary-premaxillary series 24-24; mandibular teeth
large, 24-26, fitting distinctly within the upper series.

Skin of head rather heavily pitted (in the type the epidermis has
been recently shed and this character is not obvious); a strong
nuchal fold curves across throat and from its edges arise grooves
Avhich ascend the side of neck; on the occiput the musculature causes
ridges and grooves in the skin; two prominent ridges converge on
the back part of head with a distinct median groove between them
which continues along the dorsal surface of the body and tail; two
grooves pass back irregularly to join the nuchal groove; these are
traversed by a groove which crosses throat and passes upwards be-
hind, or across, angle of mouth; eleven costal grooves, the axillary
and inguinal grooves not indicated (somewhat apparent in a para-
type).

Skin of the dorsum not strongly pitted; on the tail, the pits are
similar to those on the head; tail very slightly constricted at base;
about seven or eight vertical grooves indicated, these near the base;
tail 52 mm. long, more than a fourth longer than head and body.

Fingers, 1, 2, 4, 3, in ascending order of size, the inner greatly re-
duced and wholly included in web; web includes most of the proxi-
mal phalanx of the second and fourth and extends to and includes a
part of the second phalanx on the third digit ; terminal part of digit
very slightly spatulate; toes, 1, 5, 2, 4, 3, in ascending order of size,
the first included in web; web includes all of the proximal phalanx
and part of adjoining phalanx of the four outer digits; pads under
tips of digits prominent; limbs when adpressed overlap the width of
one fold; a small glandular spot behind insertion of femur.

Color in alcohol. Above, nearly uniform brown save that the
lower eyelid is yellowish or cream and a minute touch of lighter
color present at tip of snout and on the subnarial swellings; below,
brownish, of a very much lighter shade; chin and under hands and
feet brownish white.

Measurements, in millimeters, and data on OedipiiH multidcntata.
Type and paratypes. MCZ Nos. 14812, 14811, 14810, respectively.

Taylor: Mexican Salamanders 291

Sex ^ , ^ ,yg,.; snout to vent, 39.5, 31.5, 21.1; length of snout, 2.7,
2.3, 1.7; snout to arm, 13, 10, 1.5; axilla to groin, 20.4, 15.2, 13.5;
tail, 52, 40, 29; head width, 6.2, 5.2, 4.4; head length to jaw angle,

8.5, 7, 5; forearm, 10, 10, 7; leg, 12.1, 10, 7.1; head width into head-
body length, 6.37, 6, 5.4 times; head length into head-body length,

4.6, 4.5, 4.8 times; maxillary-premaxillary teeth, 24-24, 21-21, 23-23;
vomerine, 7-8, 6-5, 6-6; mandibular series, 24-24, , 23-22.

Variation. Practically no variation is evident save that recorded
above. The amount of overlapping in adpressed limbs is less in
smaller specimens.

Remarks. As stated, this species is most closely related to Oedipus
chiropterus, and resembles the latter rather strongly. The large
number of teeth, typical of the females of chiropterus, are present
here in the males. The males of chiropterus, in the adult condition,
ha^'e no posterior maxillary teeth, and those that are present an-
teriorly are larger than the teeth in the female. The sexual di-
morphism in chiropterus is striking as regards dentition.

Oedipus chiropterus (Cope)

18G3. Spelerpes chiropterus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1863, p. 195;
(type description ; type locality, Mirador, ^■eI■acruz, Mexico. Dr. Sartorius, collector) ;
and, 1869, p. 106; Boulenger, Cat. Batr. Grad. s. Ecaud. British Mus., 2d Ed., p. 67;
Brocchi, Mis.sion Scientifique au Mexique et dans I'Anierique Centrale, Etude sur Batraciens,
Livr. 3, 1883, p. 109; Cope, Bull. U. S. Nat. Mus., No. 32, 1887, p. 8; and No. 34, 1889,
p. 162; Duges, La Naturaleza, 1896, (2), 2, p. 482; Gunther, Biologia Centrali-.\niericana,
Rept. and Batr. 1902, p. 298; Gadow, Proc. Zool. Soc. London. 1905. p. 203, and Zoiil.
Jahrb.. 1910, p. 714.

1865. Spelerpes orculus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1865, p. 196 (type
description; type locality, Mexican tableland and southern mountains); Herrera, La Naturaleza
(2), 1, p. 340; and 1892, p. 49.

1924. Oedipus chiropterus Dunn, Field Mus. Nat. Hist., Zool. Ser., XU, 1924. pp. 99-
100; Salamanders of the Plethodontidae, Smith College 50th Ani\-. Publ., 192(1, p|i. 354,
368-371 {part, not MCZ Xos. 8404, 8408).

This small species appears to be widely distributed at high eleva-
tions in Central Mexico; in these localities it is the most common
species.

The following specimens of 0. chiropterus are in the collection:
EHT-HMS Nos. 4129-4156, 4158-95; 4197-4232, 4381-4383, 4385-
4388, 4390-4404, 4406-4430, 4432-4446, 4448-4454, .July 30, 31, 1932,
and 4460-4499, .Tuly 11, 1932, from near Rio Frio, Puebla, eleva-
tion 9,000 to 10,000 feet; Taylor and Smith.

Nos. 12144-12165, 12167-12177, 12179-12185, Aug. 12 at km. 58,
near Rio Frio, Puebla, elevation about 10,000 feet; Taylor.

Nos. 4233-4312, 4500-4503, July 18, 1932, Taylor and Smith, and
Nos. 12000-12006, 12008-12029, 12031-12039, 12041-12047, 12053,

292 The University Science Bulletin

12060, 12061, 12071, 12076, 12082, 12087, Sept. 2, Taylor, Cruz
Blanca and slopes of Cofre rle Perote up to 11,000 feet, Veracruz.

Nos. 3971-3972, Sept. 15, 1935, Taylor; No. 4380, July 11, 1932,
Taylor and Smith, and Nos. 4512-4514, 4516-4517, 4521-4532, Tay-
lor, km. 50 near Tres Cumbres. Morelos; Nos. 12271-12291, 12293-
12329, 12331-12333, 12335-12411, 12417-12419 km. 35 near Tres
Cumbres, in a pedrigal. July 6, 1936. Taylor and Smith.

Nos. 12124-12140, 12233, 12234, 12237, 12240, 12243, 12246,
12253, 12254, 12261, 12262, 12264, 12420-12428, 12430-12432, 12434-
12437, 12440-12488, 12490-12492, Lake Zempoala, Morelos, elevation
10,000 to 11.000 feet, Aug. 5, 6, 1936, Taylor.

Diagnosis. A small species with the feet and hands partially
webbed; first finger and first toe short, completely involved in the
web ; 13 costal grooves ; vomerine teeth in two very short series of six
or seven teeth tending to meet at an oblique angle, the series sepa-
rated by a distance eciual to the normal space between two teeth; ex-
tending to inner level of choanae; females with a maxillary-premax-
illary series of 20-24 teeth, the series beginning about middle of eye;
adult males with about six or seven teetli in the series beginning
much in ach'ancc of the eye, the teeth much enlarged; the four pre-
maxillary teeth much enlarged, but while visible externally do not, or
only occasionally, pierce the lij); no canthus rostralis; large, plainly
visible hedonic gland on tip of chin in male, and a small gland
behind and slightly above insertion of hind limb; a continuous
median dorsal groove not reached by the lateral grooves. Tail
longer than snout-to-vent length. Color variable; dull grey or
bluish black on entire dorsal and lateral surfaces, with ventral sur-
faces lighter; or dorsal surface creamy-vinaceous, with sides black-
ish. Sometimes the lighter cream color forms two dorsolateral lines,
the vinaceous a median stripe; others are uniformly lighter, nearly
lavender above with lighter shades below. One specimen is cream
with tlie pigment segregated to form irregular spots above and
below.

Variation. The vinaceous pattern at times is so constant that one
feels tliat one is dealing with a distinct race; yet this same variation
crops up with greater or lesser degree of frequency throughout the
range. The limbs when adpressed are separated by from four and
one half costal folds to two costal folds. The higher number usually
applies to females, while in typical males the limbs are separated
by two to two and one half folds.

Reinarks. The species is known from the states of Veracruz,

Taylor: Mexican Salamanders 293

Puebla, Mexico, Morelos and Distrito Federal. I failed to obtain
specimens of the species on Nevada de Toluca at elevations where
one would normally expect them to be plentiful.

Cope's Spelerpes orculus from the Mexican Tableland seems to be
properly associated with this species as a synonym.

Oedipus pennatiihfs (Cope)

1869. Thoriux pennatribus (typ. err.) Amer. Nat., 18G9 p. i'i^.

18fi9. Thoriiis pcniiatuluK Cope, Proc. Acad. Nat. Sci. Pliiladelphia, 1809, pp. 111-112;
(type description; tyjie locality, "Orizava." Mexico, F. Siiinichrast, collector; type UvSNM.
No. 63-tl originally); Boulenger, Cat. Batr. Grad. British Mus. (2), 1882, p. 79, pi. 3.
fig. 2 (head and neck) (Orizaba, Mexico); Cope. Bull. U. S. Nat. Mas., :U, jil. 2T, figs. 2-4
(Skull, fide, Dunn 1926); Glinther, Biologia Centrali-Americana. Rept. Batr., 1902, pp. 304-
305; Gadow, Proc. Zool. Soc. London, 1905, p. 202.

1883. Thoruis (sic) pennatulus Brocchi, Mission Scientifique au Mexi<iue et dans rAniericpie
Centrale, part 3, sec. 2, Etude des Batraciens, Livr. 3. 1883, p. 119.

1922. Oedipus pennatulus Dunn, Proc. Biol. Soc. Washington, 35, Mar. 20, 1922, p. 6;
and Field Mus. Nat. Hist., Zool. Ser., XII, May. 19, 1924. pp. 99, 100 (key); and The
Salamanders of the Family Plethodontidae, Smith College Fiftieth Anniversary Publications,
1926, pp. 469, 374-376, fig. 64, map.

?1877. Spelerpes sp. Wiedersheim, Morph. Jahrb. 3, 1877. pp. 427, 482, 498, pi. 21,
fig. 48, pi. 24, fig. 87 (Veracruz; Dunn suggests that this should be in this synonymy. If
so it is likely the locality refers to the state, not the city).

?1877. Spelerpes minimus Wiedersheim Inr rit. p. 544 ^"eracruz).

Description of species. (From EHT, Nos. 12141-12143, 12343A,
12343B; collected near Acultzingo, Veracruz, Aug. 14, 1936, by E.
H. Taylor.) Very small species; body moderately slender, the tail
about one and one third times as long as head and body; snout
blunt, somewhat oval, seen from above, the eyes extending beyond
outline of head; no canthus rostralis, and no longitudinal groove be-
hind eye; first vertical groove close behind angle of mouth; eye
large, its length greater than length of snout; both eyelids fitting
under the fold of skin behind eye; nostril very large, its diameter
more than one third of eye; edge of lip slightly swollen at the
groove; angle of the jaw a little behind the angle of the eye; width
of head contained in head-body length, 7.3 times; head length in
head-body length, 5.6 times; 13 costal grooves from axilla to groin;
the foreleg covers three costal folds, the hind leg four; when limbs
are adpressed, separated by 6 folds; limbs weak, fingers 3, 2. 4, 1
in descending order of length; one and four are not free at tip; toes
3, 4, 2, 5, 1 in descending order of length, the two outer toes rudi-
mentary, not free at their tips; anal lips slightly rough; vomerine
teeth in short series not extending beyond nares; three to five teeth
in series; no teeth on maxilla; parasphenoid teeth in a single, rather
broad patch; three teeth on the premaxilla.

Color. Above brown, color only slightly lighter than the dark
lateral bands which merge imperceptibly into the ventral colora-

294

The University Science Bulletin

tion (under magnification the ventral coloration consists of closely-
set circular, cream dots separated by black reticulation) ; a few
lighter flecks on ventral surface of belly and chin.

Variation. The dorsal lighter band is evident on all the speci-
mens and the light flecking is present on ventral surfaces and sides;
the head is the color of the sides; one specimen has a suggestion of
a dim row of darker flecks on the dorsal median line.

Measurements and data on Oedipus pcnnatuliis Cope

Number

Snout to vent

Snout to gular foLi, below

Snout to foreleg

Tail

Width of head

Arm

Leg

Axilla to groin

Costal grooves

Grooves on tail

12141

25.2
4.5
7.1

33.2
3.45
3.4
4

15.4

13

40

12142

18.4
3.6
5.2

19*
2.8
3.2
3.3

10.8

13

28*

12143

21.2

4

6.1
28

3.5

3.5

4
12
13
39

2245

20.2
3.8
6.1

25
3.8
3.8
4.1

12.3

13

32 +

2244

21.2
4
6.5

22.5*
3.5
3.5
4.1

13.5

13

22 +

* Not complete.

Remarks. These small specimens were taken under rocks at the
top of a mountain south of Acultzingo (where the highway crosses).
When found they were closely coiled in a watch-spring spiral, and
remained so coiled until about five seconds after being placed in
alcohol.

Oedipus lineolus (Cope)

(PI. XXIX, fig. 3)

1865. Spelerpes lineolus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1805, p. 197 (type
locality, Mexican tableland; C. Sartorius, collector); idem, 1866, p. 132 (Orizaba, Vera-
cruz); idem, 1868, p. 313 (Cordova, Veracruz); Bouleuger, Cat. Batr. Grad. British Mus.,
(2) 1882, p. 74 (Orizaba, "Mexico"); Brocchi, Mission Scientifique an Mexique et dans
TAmerique Centrale, Etude des Batracjens, part 3, sec. 2, livr. 3, 1883. p. Ill; "Orizava"
Veracruz, Giinther, Biologia Centrali-.\mericana, Batrachia. Jan., 1902, p. 304 (Orizaba).

1869. Opheoatrachus lineoius Cope, Proc. Acad. Nat. Sci. Philadelphia, 1869, pp. 101-102
(Eastern Mexico).

1884. Geotriton lineolus Garnian, Bull. Es.sex Inst. 16, 1884, p. 39.

1887. Oedipina lineolus Cope. Hull. U. S. Nat. Mus., 32, 1887, p. 8.

1896. Oedipina lineola Cope. A\wv. Nat., 1896, p. 1022 (distribiition).

1924. Oedipus lineolus Dunn. Field Mus.. Nat. Hist. Zofil. Ser., XII. No. 7, Pulil. 221.
May 19, 1924, pp. 99, 100; and The Salamanders of the Family Plethodonf idae. Smith College
Ann. Ser., 1926, pp. 422-425, map, fig. 81 (Mexican tableland ; Jalapa, Veracruz).

?1879. Spelerpes (.Oedipus) iiifuscatus Peters, Monatsb. Konigl. preuss. Akad. Wiss. Ber-
lin, 1879, p. 778 ("Hayti." Seems probable that this locality is erroneous).

?1902. Spelerpes uniformis (part.) Giinther, Bioligia Centrali-Americana, 1902, p. 304.

Taylor: Mexican Salamanders 295

Description of the species. From EHT-HMS, No. 2415; collected
about 10 km. southeast of Cordova, near San Lorenzo, Veracruz,
Aug. 20, 1987.

Body slender, cylindrical, wornddvc; the tail elongated, nearly as
thick as body, tapering rather suddenly near the tip; snout blunt,
truncate, the eyes protruding beyond outline of head when seen
from above; no canthus rostralis; no longitudinal groove behind eye;
vertical groove a little behind angle of mouth; visible below, and
arising to upper level of eye; angle of jaw is much behind posterior
angle of eye ; both eyelids fit under a fold of skin posteriorly ; second
groove, forming a nuchal fold below, and practically encircling neck;
eye distinctly longer than the snout ; nostrils small ; no swollen area
about the narial groove; 14 costal grooves, counting the dim ones
in axilla and groin; about 43 grooves on tail. Limbs very slender,
short, the anterior reaching to about the third costal groove, the
hind leg reaching to very near 12th groove, leaving the adpressed
limbs separated by 9 complete costal folds; fingers very minute,
their descending order of size, 3. 2, 4, 1, the tip of the first involved
in skin, others free; order of length in toes, 3, 2, 4, 5, 1, the tip of
first toe not free; anal lips apparently smooth.

Vomerine teeth, in two curved series, consisting of 10 and 13
teeth, meeting medially; parasphenoid teeth in two groups, con-
fluent anteriorly, divergent posteriorly, separated from the vomerine
teeth by an interval more than half distance between the choanae;
maxillary and premaxillary with teeth.

Color. Above dark, nearly black; below grayish-black (under
magnification the ventral coloration consists of minute round cream
dots of various sizes, separated by black). A few scattered cream
flecks on the chin, throat and side of head, especially below eye.

Measurements iin mm.). Snout to vent, 34.5; snout to arm, 9.4;
width of liead, 3; axilla to groin, 21.2; tail, 51.5; total length, 86;
head width, in snout-to-vcnt length, 11 times. (I suspect that the
tail has been reproduced. )

Remarks. I found this specimen on the side of a rocky hill south-
east of Cordova. The specimen was hidden in a pile of wet chips
about the base of a stump.

The species has remained very rare in collections. Dunn (1926)
was able to examine but three specimens. From the measurements
recorded, either the species is variable or the differences in propor-
tion are due to age and sex. Dunn (1926) has referred the Haitian,
"infuscatus" of Peters, to this form.

296 The University Science Bulletin

Oedipus salvinii Gray

(Plate XXVIII, fig. 2)

1868. Oedipus salv,„!i Gray, Ann. Mag. Nat. Hi.st. (4), 2, j,. i'.n (tvjie ■lesciiption •
t.vpe locality, Guatemala, Pacific Coast; O. Salvin. collector); Sumichrast, Bull Soc Zool
France, 5, 1880, p. 190; and La Natwaleza, G, 1882, p. 79; Dunn. Field Mus Nat Hist
Zool. Ser., XII, May. 1924, pp. 99, 100 (key); The Salamanders of the Family Pletho-
dontidae. Smith College Fiftieth Anniversary Publications. 1926, pp. 4(i5-4ns fi^ ' 74 map
Schmidt, Field Mus. Nat. Hist., Zoiil. Ser., XX, No. 17, Oct. 31, 1936, ],p. 147-14,s, fig. 17.

1879. Oedipus carboiiarim salvinii Cope. Proc. Amer. Philos. Soc, 18, 1879, p. 267*!

1882. Sperlerpes varinjatus Boulonger, Cat. Batr. Grad. s. Caud. Briti-h Mus 'W (d
1882. p. 73 (part.).

1887. Oedipus variegatus salvinii Cope, Bull. U. S. Nat. Mus., No. 32, 1887, p. 8.

1878. Spflcrpes sa/viuii Miiller, Verh. Naturf. Ges. Basel, 6, 1878, ,.. .",79 ("Guatemala");
Strauch, Salamanders, p. 84 (not .seen); Brocchi, Mission Scientifiqur an Mexique et daiw
L'Amenque Centrale, Etude sur Batr., livr. 3, 1883, p. 117, pi. 18, figs. 3, 3a. 3b, 4. 4a, 46.
(Tehuantepec); De Leon, Indice de los Batracios que se encuentran tn la Republica' .Me.\icana,
Tacahuya, .June, 1904, p. 38.

1896. Sprlerpes varieuatus Werner. Verh. Ges. Wien, 46. p. 351 ; Giinther, Biologia
Centrali-Aniericana, Kept, and Batr., .Jan.. I!i02. p. 302 (part.). \,l. 7"), fig. D.

1878. Spelerpes .sp. affin. sp. .'<alvniii Miiller, Verh. Natun. Ges. Basel, 6, 1878. pp. 579,
645, pi. III. figs. C. C, D (Guatemala).

Description of species. (From EHT-HMS, No. 3995, 5 ; col-
lected at Tonola, Aug. 27-31, 1935.) (Taken in a freioht car on
railway.) Body, typical salamander form. Head flattened, the
outline a truncate oval, distinctly wider than body; eye relatively
small, its length not as long as snout, but about equal to its distance
from the nostril; the hind part of eyelids inserted under a fold of
skin; nostrils small; a swelling IjcIow nostril al)out groove on upper
lip; the angle of the jaw extends far behind the posterior corner of
eye. An ill-defined shallow groove from eye to first vertical groove.
A strongly-defined gular fold which reaches up only a short distance
on the sides of neck; vomerine teeth in two curved series, practically
meeting medially, and extending laterally beyond the choanae;
about 15-17 teeth in a series; parasphenoid teeth in a large patch
pointed anteriorly, posteriorly with a median notch, separated from
the vomerine teeth by twice width of a choana; maxillary tooth
series large, extending back to the middle of the eye socket; thirteen
costal grooves; the adpressed limbs separated by three and one half
folds; limbs strong, the arm reaching eye; fingers fully webbed,
flattened; the front outline of hand scalloped; toes completely
webbed, flattened. Tail sliglitly longer than head and body, dis-
tinctly constricted at base; anal lips smooth or slightly folded.

Color. Above deep purplish lavender broken into irregular areas
and surrounded by cream borders; beginning on snout an irregular,
yellow, more or less discontinuous line extends dorsolaterally ; be-
ginning behind the eye is a broad chocolate band, very irregular on
its upper edge, ami fairly straight on lower edge (somewhat laven-

Taylor: Mexican Salamanders 297

dcr on the upper edge). Lips and ventral surface of head and body
cream-yellow with a few flecks of darker color; under tail same,
but flecked more heavily with lavender; tail dark puri)lish lavender,
reticulated on at least proximal lialf with cream.

Measurements {in mm.). Snout to vent, 81; snout to jaw angle,
10.2; snout to gular fold (ventral), IJ); snout to insertion of arm,
24; width of head, 12; tail. <S(r, arm, 18; leg, 19; axilla to groin, 50;
width of head in snout-to-vent measurement, 6.75 times; head length
into snout-to-vent measurement, 4.26.

Remarks. Since this specimen was taken in a freight car used
for transporting bananas, its provenance is uncertain; however, it
seems very probable that it originated on the coast, most probably
in southern or central Chiapas.

Oedipus platydactylus (Cuvier)

(Plate XXVUL fig. D

1831. Salaniaridra variegata Gray, in Griffiths Cuvier's Animal Kingdom, 9, p. 107 (not
of Bory St. Vincent, 1829, Diet. Class. Hist. Nat., 15, p. 08) (type description; type
locality, Mexico).

1831. SalaiiKiiidra plat ydactyltis Cmier, in Graw CiriRifh's Cin-i;'r's Animal Kingiloin, ii.
107.

1838. Oedipus plati/dactyliis Tschudi, Mem. Soc. Sci. Nat. Neuchatel, 1838, p. 58; Dunn,
Field. Mus. Nat. Hist. Zool. Ser., XH, May, 1924, pp. 99, 100; and The Salamanders of the
Family Plethodc.ntidae, Smith College, Fiftieth Aniii\-ersary Publications. 1920, p|). 400-403,
fig. 73, map.

1850. Oedipus rariegatus Ciray, Cat. Batr. Grad. British Mus., 1850, p. 48; Ann. Mag.
Nat. Hist., 2, 1858, p. 300; Moore, Proc. Acad. Nat. Sci. Philadelphia, 1900, p. 019;
Fowler and Dunn, Proc. Acad. Nat. Sci. Philadelphia. 1917, p. 19.

1854. Bolitoglossa Mexicana Dumeril and Bibron (part.). Erp. Gen., \' . 9. 1854, p. 93,
pi., 104, fig. 1 (Dolores, Peten, Guatemala).

1800. Geotriton carbonarius C-ope, Proc. Acad. Nat. Sci. Philadelphia, isiio, \>. 373;
(type description; type locality, Jalapa, Mexico); .Journ. Acad. Nat. Sci. Philadel|ihia (2),
6, 1866, p. 98.

1809. Oedipus carbonarius Cope, Proc. ,\cad. Nat. Sci. Philadel|]hia, 1809, |i. 103; and
Proc. Amer. Philos. Soc, 1879, 18, p. 207; Sumichrast, Bull. Soc. Zoiil. France, ISSl, ji.
231; and La Naturaleza, 6, 1882, p. 78.

1870. Geotriton variegata Garman. Bull. Essex lust., 10, 1884, \). 39.

1883. Spelcrpes Mexicaimm Brocchi, Mi.ssion Scientifique au Mexique, Etuile sur les
Batraciens, livr. 3, 1883, p. 113, pi. 18b, figs. 1, 2, 3, 4.

1883. SpeJerpes Copei Brocchi, loc. rit.. p. 113.

1883. Spelerpe.s punctatum Brocchi, lor. rit.. p. 115, jil. 20, figs. 345 (tj'pe description;
type locality, .\lta ^'era Paz, Guatemala).

1870. Spelerpes variegatus Strauch, Mem. .\cad. Sci. St. Petershourg (7), 16, 4, p. 84;
Miiller, ^'erh. Naturf. Ges. Basel, 1878, 0. ji. 579; Boulenger, Cat. Batr. CJrad. s. caud.,
2d ed. 1882, p. 73; Ixlnnberg, Zotil. .\nz., 1899, p. 545; Glinther, Biologia Centrali-.-\meri-
cana, Batr. Rept., Jan., 1902, pp. 302-303, pi. 75, fig.s. A. B. C. ; Werner, Abh. Bayer .\kad.,
22, 1903, p. 352; Gadow, Proc.^Zool. Soc. London.. 1905, p. 203; and Zool. ,Iahrb., 1910. p.
305; Ruthven, Zool. .Jahrli., 23. 1912. p. 305; and Rei)t. Michigan .\cad. .S'i.. 14, 1912.
p. 231.

1884. Geotriton laregata Garman. Hull. Essex Inst., 10, 1884, p. 39.
1884. Geotriton Mexicana Garman. Bull. Essex Inst. 10, 1884, p. 40.
1894. Spelerpes Mexicana Duges, La Naturaleza (2), 2, 1894, p. 377.

298 The University Science Bulletin

The collection contains specimens of this species from the follow-
ing localities: EHT-HAIS, Nos. 3964, 15 mi. S. Valles, San Luis
Potosi. Jmie 13, 1932, E. H. Taylor and Hobart M. Smith, col-
lectors; 15200, near San Lorenzo, Cordova, Veracruz, Aug. 19, 1936,
E. H. Taylor, collector; 15201, Potrero Viejo, Veracruz, Aug. 23,
1937, E. H. Taylor, collector; 15202, 5 miles each of Cordova, Vera-
cruz, H. R. Roberts, collector.

Description of species. (From EHT-HMS, No. 15202.) A large
Oedipus, the body moderately robust. Head seen from above, trun-
cate oval, the eyes moderately prominent, extending slightly beyond
outline of head; nostrils small, with a prominent swelling below
them near the lip about the groove; length of eye slightly greater
than its distance to nostril, but longer than the snout; a groove from
behind eye joins the first nuchal groove, which is continuous across
the throat, though but dimly visible (strongly visible in No. 15200) ;
second nuchal groove crosses neck and arises high on the side of
neck; angle of jaw^ far back of posterior corner of the eye; posterior
parts of eyelids fit under a fold of skin; body with thirteen costal
grooves from axilla to groin; the limbs, where adpressed, separated
by about four folds; limbs strong, both fingers and toes well de-
veloped, flattened, enclosed in webs; the anterior edges of the pal-
mate hands and feet scalloped; anal slit without papillae; but the
inner walls strongly folded; tail tapering gradually, slightly com-
pressed.

Vomerine teeth in two curved series of 13 teeth each, almost in
contact medially, extending to outer (lateral) edge of the choanae;
a single large patch of parasphenoid teeth, separated from the
vomerine teeth by the width of a choana.

Color. Above the color is light buff to fawn, and covers the whole
dorsal surface, save head, which is more or less covered by a dark
triangular patch, the apex of which extends back on the neck; a few
lighter fiecks on the head; sides of head, body and tail and all ven-
tral surfaces dark plumbeous to blackish, practically uniform.

Measurements of Oedipcs platydactylus (Cuvier)

No. 15202 No. 1.5200

Snout to vent 72 43

Snout to jaw angle 9 6.8

Snout to fiular fold 16.2 10.3

Snout to forelimb 21 13

Width of head 10 7

Axilla to groin 43 . 5 26 . 6

Arm 13 9

Leg 15 10

Tail 73 42

Taylor: Mexican Salamanders 299

Variation. Nop. 15200 and 3964 have some black flecks or streaks
on the back and taiL I recently examined a large specimen from
Tamazunchale, San Luis Potosi. belonging to Mr. Ottys Sanders,
Dallas, Texas, in which the lateral and ventral coloration was coal-
black.

Remarks. Certain of the more striking color variants have been
described as distinct species {variegatiis, carbonarius) ; it seems
likely that Dunn is correct in placing these in synonymy. How-
ever, the buff-colored form here discussed is apparently the typical
platydactylus.

The specimens from San Luis Potosi carry the range much farther
north than known heretofore.

Gymnopis multipUcata oaxacae Mertens

1930. Gymnopis multipUcata oaxarae Mertens, Abh. Ber. Mus. Natiir. — Heimatk. Natur.
Ver. Magdeburg, Bd. VI, heft II, 1930, pp. 153-1,55, fig. 14 (from Blatter Aquar. Terr-Kiinde
1928); (type description; type locality, Cafetal Concordia, 600 m. between Puerto Angel
and Salina Cruz, Oaxaca, Mexico; Lafrentz, collector).

A single specimen, EHT-HMS, No. 4604, of this rare form was
collected by H. M. Smith on the hills east of Tonola, Chiapas. It
presents the following characteristics:

A total of 131 primary folds anterior to anus; the 13 anterior
folds and the posterior 18 completely encircle the body — the total
of primary and secondary folds on body and tail, 236; a strongly
developed nuchal fold with lateral grooves meeting medially; first
nuchal groove crosses throat, passes back of the angle of jaw and
across the head; the area between the two aforementioned grooves
partially divided by a short transverse groove on the dorsal part of
the neck, and by a somewhat longer transverse groove on the throat;
nostrils small, the distance between them (2.5 mm.) less than the
distance between the nostril and the globular tentacle (3.2 mm.) ;
eye to nostril (4.6 mm.) a little less than interorbital distance (5
mm.), which equals the length of snout; snout projecting 1.7 milli-
meters beyond mouth; 15-16 maxillary teeth on each side, inner,
vomero-palatine series 16-18; a single series of teeth in lower jaw,
11-12 on each side, large, conical, much larger than teeth on upper
jaw; two small papillae in the preanal region; four or five in the
postanal region; a prominent lateral "fold" or ''ridge" extends the
length of tlie body.

Measurements {in mm.). Length of head to angle of the jaws,
11.1; width of head, 7.3; snout to gular fold, 12.5; total length, 340;
tail. 2; Ixxly width, 10; body width in length, 34 times.

300 The University Science Bulletin

Remarks. It may be noted that this form departs from the typi-
cal generic characteristics in hicking the inner row of teeth in lower
jaw; the teeth in general, instead of being small and siibequal, are
relatively large and vary greatly in size in the lower and iipjier jaws.

The specimen was found under a log burrowed in hard earth near
to the top of the large hill about three miles east of Tonola.

I acknowledge indebtedness to Dr. E. R. Dunn for the identifica-
tion of this species under the name here used.

LITERATURE

1893. Blatchley, AV. S. On a collection of Batrachians and Reptiles from
Mt. Orizaba, Mexico, with descriptions of two new species. Proc. U. S. Nat.
Mus. XVI, 1893, pp. 37-42.

1882. BouLi-^xcEK, G. A. Catalogue of the Batrachia Gradiontia s. Caudata
and Batrachia Ajioda in the collection of the British Museum, 2d Ed., 1882,, pp.
1-118. pis. 1-9.

1881-1883. Brocchi. Mi.^sion Scientifique au Mexique et dans I'Amerique
Centrale. Etude des Batracien.s livr. 1-3. 1881-1883, pis. 1-21.

1865. Cope, E. D. Third contribution to the Herpetolojiy of Tropical
America. Proc. Acad. Nat. Sci. Philadelphia, 1865, pp. 185-199.

1867. A review of the species of the Amblvstomidae. Proc. Acad.

Nat. Sci. Philadelphia, Dec, 1867, pp. 166-211.

1869. A review of the species of the Plethodontidae and Desmog-

nathidae. Proc. Acad. Nat. Sci. Philadelphia, May. 1869, pp. 93-118.

1889. The Batrachia of North America. Bull. U. S. Nat. Mus.; No.

34, 1889, pp. 1-525. Plates 1-LXXX\T; text figs. 1-119.

1870. Denies, Alfredo. Una nue\a especies de ajolote (achociue de agua)
de la laguna Patzcuaro. La Naturaleza, I. 1869-1870 (1870). p. 241. Plate 5A.

1896. _ Ambvstoma altamirani A. Duges. La Naturaleza (2), IL 1896,

pp. 459-461, pi. XXIX.

1854. DuMERiL, A. M. C.. and Bibron, G. Erpetologie generate de histoire
naturelle complete des Reptiles. Vol. IX, 1854, pp. 1-XX; 1-440.

1924. Di'NN, E. R. New salamanders of the genus Oedipus, with a svnop-
tical key. Field Mus. Nat. Hist., Zool. Ser., NIL 1924, pi). 95-100.

1926. The salamanders of the family Plethodontidae. Smith College

fiftieth anni\ersarv publications VII, Northampton, Mass., pp. l-VIII ; pp.
1-441; tigs. 1-11. Maps 1-86. 1 plate.

1928. A new genus of .salamanders from Mexico. Rhyacosiredon gen.

nov. Proc. New England. Biol. Club, X, No. 13. \)\). 85-86.

1926. G.KDOw, H. Der Mexikanische Axolotl. Blatter Aciuar-Tcrrar. Kunde,
37, 1926, ]). 252.

1900-1902. Gf-NTHER, A. Biologia Centrali-Americana. Batr. Feb., 1900, to
May, 1902, pp. 197-309, pis. 60-76.

1936. Herre, Wolf. Ueber Rasse und Arlbildmm. Studien an Salamandriden.
Abh.. Ber. Mus. Natur. Vorges. Natur. Ver. Magdeburg, Bd. VI. Heft. 3. 1936.
PI). 193-221.

1928. L.^FRENTz, K. Neue Beol)achtuniien an amphibien des Mexikanischen
Hochlandes. Bliitter Aquar. Terrar. Kunde, 39, 1928, pp. 89-92, 110-115, 5 figs.

1930. Ein neuer jilethodont-salamander aus Mexico. Alih. Ber. Mus.

Natur-Heimatk Natur. Ver. Magdeburg, Bd. VI, Heft. 2, 1930, pp. 150-152.

1930. Lafrentz, Dr. D. I^ntersuchen uber cUe Lebensgeschichte mexikanis-
cher Ambvstoma-Arten; Abh. Ber. Mus. Natur .-Heimatk. Naturw. Ver. Madge-
burg Bd. VI, Heft. II. 1930, pp. 91-127. 3 figs., pis. II and III.

Taylor: Mexican Salamanders 301

1930. Mertens, Robekt. Beniorkungt'ii iiher die xon Hcnn Dr. K. J.afrcutz
in Moxiko gesammelten amphibian unci Reptilien. Al)h. Bpr. Mu.-^. Natur.-
Hoiniatk. Xaturw. Ver. Magdeburg, M. VI. Hcff. 2. 1930. pp. 153-162.

1921. Noble, G. K. Anterior cranial (■kincnt.< of Oc<li/>ii.'< and cortain other
.sdaniander.-^. Bull. Amer. Mus. Nat. Hi.-^t., XLIV. 1921, pp. l-fi. pis. 1-2.

1936 Schmidt, K.\rl P. Guatemalan Salamanders of the mnus Oedii>u.-^.
Field, Mm. Nat. Hist., Zool. Scr., XX, Oct. 31, 1936, pp. 136-166, tig.^. 13-19.

1879. ^"EL.ASC0, J. M. Description metamorphosis y costumbres de una
especie nue\a del genero Siredon, encontrado en el lago de Santa Isabel, La
Naturaleza. IV, 1879, pp. 209-233. pis. VII, MIL

1930. WoLTERSTORFF, Dr. W. Zur Systematik unil Biologie der Urodelen
Mexikos. Abh. Ber. Mu.'^. Natur-Heimatk. Naturw. Ver. Madgeburg, Bd. "\'I,
Heft. 2, 1930, pp. 129-149, figs. 1-13.

302 The University Science Bulletin

PLATE XXIV

Fig. 1. Ambijstomo sp. Shaw. EHT-HMS. No. 3997. Rancho Guadalupe,
noar Sm Martin, Mexico; actual snout-to-vent measurement, 80 mm.; total
length. 150 mm. Not discussed in text.

Fig. 2. Ryacosiirdon nltamirntn (Duges). EHT-HMS. No. 12511. Near
Lake Zemiioala. Morelos, 11,000 ft. ele\ation. Actual snout-to-vent lenjith,
68.2 mm.; total length. 143.2 mm.

Taylor: Mexican Salamaxdkhs

303

PLATE XXIV

\

304 The University Science Bulletin

PLATE XXV

Fig. 1. Oedipus orizabeiisis (Bkitchley). EHT-HMS. No. 12067. Lake
Zempoala, Morelos; actual 8nout-to-vent. length, 53.5 mm.; total length. 99.5.

Fig. 2. Oedipus orizahcnsis (Bhitchley). EHT-HMS. No. 12239 (same
locality as 12067); actual snout-to-\ent length, 56 mm.; total length, 105 mm.

Fig. 3. Oedipus altamo)iin})tis sp. nov. f^HT-HMS. No. 12245. Tyiie.
Near Lake Zempoala, Morelos; 10,500 feet; actual snout-to-vent length,
48.6 mm.

Fig. 4. Oedipus altamontanus sp. nov. EHT-HMS. No. 12239. Paratype.
Same locality; actual .snout-to-vent length. 40 mm.

Fig. 5. Oedipus smilhi sp. nov. EHT-HMS. No. 3965 i . Paratype. Cerro
de San Luis. Oaxaca, Oaxaca; actual snout-to-vont length, 69 mm.; total
length, 139 mm.

Fig. 6. Oedipus smilhi s^). now EHT-HMS. No. 3966 9 . Type, Cerro de
San Luis, Oaxaca, Oaxaca; actual snout-to-vent length, 69 mm.; total length.
145 mm.

Taylor: Mexican Salamanders

305

PLATE XXV

T^

3.

20—4141

30() The University Science Bulletin

PLATE XXVI

Fig. 1. Ambyst077ia schmidti sp. nov. EHT-HMS. No. 3999. Type.
Rancho Guadalupe, 10 mi. E. San Martin (Asuncion), Mexico; actual snout-
to-\ent length, 52 mm.; total length, 89 mm.

Fig. 2. Oedipus robcrlsi sp. nov. EHT-HMS. Xo. 12503. Nevada de
Toluca, Mexico, between 10,000 and 11,000 feet; actual snout-to-vent length, 51
mm.; total length, 95.8.

Fig. 3. Oedipus cephalicns (Cope). EHT-HMS. No. 4539 $ : east of Rio
Frio, Puebla, Mexico. Actual length snout to posterior part of vent, 49.5 mm.;
total length, 98 mm.

Fig. 4. Oedipus cephalicus (Cope). EHT-HMS. No. 4536 9 ; east of Rio
Frio, Puebla, Mexico. Actual snout-to-vent length, 56.5 mm.; total length,
110 mm.

Taylor: Mexican Salamanders

307

Pl.ATK XXVI

3.

308 The University Science Bulletin

PLATE XXVII

Fig. 1. Ovdipus bcUit (Gray). MCZ. Xo. 3935, Guerrero, Hidalgo, Mexico.
Actual head-body length. 103 mm.

Fig. 2. Oedipus bellii {Gray). MCZ. No. 3938, Guerrero. Hidalgo, Mexico.
Head-body length, 53 mm.

Fig. 3. Oedipus giganteus sp. nov. MCZ. Xo. 843-4, Julapa, Veracruz,
Mexico. Head-body length, 46 mm.

Fig. 4. Oedipu.^ giganttus ^p. nov. Type. MCZ. Xo. 8435, Jalapa, Vera-
cruz. Head-bodv length, 110 mm.

Taylor: Mexican Salamanders

309

PLATE XXVII

310 The University Science Bulletin

PLATE XXVm

Fig. \. Oedipus plotydactylus (Cuvici). EHT-HMS. Xo. 15202, 5 miles
oast of Cordova, Voracruz, Mexico. Head-body length, 72 mm.

Fig. 2. Oedipus salvinii Gray. EHT-HMS. No. 3995 ?,(?) Tonola Chiapas.
Mexico. Head-body length, Si mm.

Taylor: Mexican Salamanders
PLATE XXVIII

:ni

312 The University Science Bulletin

PLATE XXIX

Fig. 1. Oedipus midtidentata sp. nov. Type. MCZ. No. 14812, Alvarez,
San Luis Potosi, Mexico. Head-body len<jth. 39.5 mm.

Fig. 2. Oedipus Icprosus (Cope). MCZ. Xo. 7659, Zometla, Orizaba,
Mexico. Head-body length, 57 mm.

Fig. 3. Oedipus lineolus (Cope). EHT-HMS. No. 2415. Near Cordova,
Veracruz, Mexico. Total length, 86 mm.

Fig. 4. Oedipus mntini sp. nov. MCZ. No. 3916. Guerrero, Hidalgo,
Mexico. Head-body length, 48 mm.

Fig. 5. Oedipus manni sp. nov. MCZ. No. 3915. Type. Guerrero, Hidalgo,
Mexico. Head-body length, 53.2 mm.

Fig. 6. Oedipus 7)iaimi sp. no\. MCZ. No. 3925, Guerrero, Hidalgo, Mexico.
Head-body length. 46 mm. Ventral view.

PLATE XX TX

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 15

Notes on the Mexican Snakes of the Genus Leptodeira.
with a Proposal of a New Snake Genus, Pseudolepto-
deira

EDWARD H. TAYLOR,

Department (if Zoology, University of Kansas

Ab.'^tract: The woik is based on si)ecimen.s of Leptodeira in the collections
made in Mexico bj- Dr. Hobart Muir Smith and the author. The following
Mexican .species are recognized as l)elonging to the genus: Leptodeira punctata
(Peters), L. spletidida Giinther, L. nn/stacitia Cope. L. dinickrri Werner, L.
scptentrionalis (Kennicott), L. jri ,iata (Cope), L. mactdata (Hallowell), L.
yucatanensis yucatanensis Cope, L. yueaiitauetisis }7ialleisi Dtinn and Stuart;
L. bressovi sp. nov. (type locality. El Sabino, Uruapan, Michoacan), L. .smilhi
s]). nov. (tyiie locality. El Sabino, Uruapan, Michoacan).

L. lalifasciata (Giinther) and L. disridor (Giinther) are placed in a new
genus Pseudoleptodeira, while the genus Hypsiglena Cope sensu strict u (in-
cluding the .species torquata, ochrorhyuchua and ajflnis) is not regarded as be-
ing a synonym of the genus Leptodeira.

AN interpretation of the species of the genus Leptodeira Fitzinger,
acceptable to herpetologists in general, appears to be difficult
to formulate; at least previous studies by numerous authors have
failed to bring about anything approaching a unanimity of opinion.
That workers have differed in their treatment of the genus may be
due, in some cases, to inadecjuate material available for study; in
other cases, to a different concept of genera and species, or to the
individual's technique of study. It is significant that, with prac-
tically the same specimens before them, such experienced workers
as Giinther and Boulenger should have arrived at such diverse re-
sults when dealing with this genus. While a complete review of the
literature is impractical here, it is pertinent that a few, more im-
portant contributions be briefly reviewed.

Giinther (1858) studied the American specimens of this genus in
the British Museum (64 specimens) and placed all in a single spe-

(31.5)

316 The University Science Bulletin

C'ies, Leptodeira annulata (Linne). These specimens were Mexican,
Central American, and South American. Later Giinther (1895) and
Boiilcngcr (18961 recognized four species in this same lot of material.

Giinther (1895) reviewed the Mexican and Central American
forms in the British Museum in the Biologia Centrali-Americana,
May, 1895, pp. 168-174, pis. LIII, fig. B, LIV, figs. A, B, and C,
LV, figs. A, B. Eight known species were recognized as follows:
Leptodeira nigrojasciata Giinther (1868); L. pacifica Cope (1869);
L. 77iystacina Cope (1869); L. annidatus (Linne) (1754); L. yuca-
tanensis Cope (1887) ; L. person({ta Cope (1869) ; L. frenata (Cope)
(1887) ; L. rhombifcra Giinther (1872). Four new species were de-
scribed, and three were figured in this same work: L. affinis, L.
splendida, L. poh/sticta, and L. ocellata.

Kennicott's species, Dipsas septentrionalis, was placed in the syn-
onymy of L. annulata (Linne) ; and Peters' Crotaphopeltis punctata
was not recognized as a Mexican form. Cope's Sibon septentrionale
rubricatum, Lacepede's Coluber albofusca, and Hallowell's Mega-
lops maculatus were not allocated.

It has been impossible to determine exactly the number of speci-
mens available to Giinther, but the year following the publication of
his work there were 79 specimens from Mexico and Central America,
and 26 from South America in the British Museum, as listed by
Boulenger.

Boulenger, in 1896, reviewed the entire genus, having at hand the
same and probably no more specimens than were available to Giin-
ther. His conclusions were strikingly different. He recognized in
the Mexican-Central American region only seven species, as follows:

(1) Leptodeira punctata (Peters), which included L. pacifica Cope;

(2) L. nigrojasciata Giinther, including L. mystacina Cope; (3) L.
frenata (Cope); (4) L. septentrionalis (Kennicott) ; (5) L. per-
sonata Cope, in which were included L. rhombifera Giinther and L.
splendida Giinther; (6) L. ocellata Giinther, in which was placed
Sibon septentrionale rubricatum Cope (with a ciuestion) ; (7) L.
albofusca (Lacepede), with which were synonymized L. yucatanensis
Cope, L. polysticta Giinther and L. affinis Giinther. L. annulata
(Linne) was restricted to South America.

Cope (1900) published a key to the genus, in which he listed seven
species. This was virtually a reprint of a key published earlier
(1891), in which the same forms were recognized. He synonymized
his L. mystacina with L. nigrofasciata. and lists L. pacifica, L.
frenata, L. personata, L. yucatanensis, L. septentrionalis and L.

Taylor: Notes on the Genus Leptodeira 317

annidata. It seems likely that he had not considered either Giin-
ther (1895 » or Boulenger (189()) in fabricating the key published
in 1900.

Mocquard (1908) treated of Mexican raid Central American spe-
cies in the Paris Museum. He accepted the disposition of species
and their synonymies made by Boulenger, and adds another species,
Leptodeira guilleni, described by Boulenger in 1905.

Werner (1913), in a key to species of Leptodeira, recognizes L.
dunckeri, L. punctata, L. nigrofasciata, L. frenata, L. scptentrionalis,
L. personata, L. gidllcni, L. annidata, L. occUata, and L. albofusca.

Perhaps the most extraordinary treatment is that of Amaral
(1929). He places all species of New World Leptodeira in a single
species, L. annidata, and recognizes four subspecies, L. a. septen-
trionalis, L. a. punctata (with nigrofasciata in synonymy!), L. a.
personata (with L. frenata and L. guilleni as synonyms!) ; L. a.
a)inid(da (includes maculatus, albofusca, dunckeri, weiseri as syn-
onyms).

Dunn (1936) published some notes on North American Leptodeira
proposing certain changes in the concept of the genus. He had
available 957 specimens, of which 606 were referred to L. rhombifera.
Of the total 400 consisted solely of heads.

Of Leptodeira {sensu strictu), Dunn recognized the following
forms: (1) Leptodeira annidata annidata (including affinis Giin-
ther) ; (2) L. annidata polysticta; (3) L. rhombifera Giinther (in-
cluding rubricatnm Cope, splendida Giinther, and ocellata Giinther) ;
(4) L. pacifica Cope (placing Crotaphopeltis punctata Peters as a
questioned synonym); (5) L. yucatanensis yucatanesis (Cope); (6)
L. yucatanensis malleisi Dunn and Stuart; (7) L. frenata Cope; (8)
L. septentrionalis septentrionalis (Kennicott) ; (9) L. septentrionalis
maculata (Hallowell) (including personata Cope) ; (10) L. mysta-
cina Cope; (11) L. nigrofasciata Giinther.

Aside from these forms the following are included in the genus:
Hypsiglena torquata, as Leptodeira torcjuata torquata Giinther;
Hypsiglena ochrorhynchus Cope as Leptodeira torquata ochrorhyn-
ckus; and Hypsiglena venusta (Mocquard) as Leptodeira tor-
quata venusta; Hypsiglena discolor Giinther as Leptodeira discolor
and Hypsiglena latifctsciata Giinther as Leptodeira latif asciata , in-
cluding as a synonym Leptodeira guilleni Boulenger.

Comments on these various proposals are discussed under the
heading of the genus or indiA'idual species.

Since 1932 Dr. Hobart M. Smith and T have been segregating a

318 The University Science Bulletin

collection of Mexican ampliibians and reptiles, among which are
some 70 specimens belonging to the genera Leptodeira and Hypsi-
glena.

Due to the courtesy of Dr. Leonhard Stejneger and Dr. Doris
Cochran I have recently examined, in some detail, Mexican speci-
mens of Leptodeira in the United States National Museum and a
number of Central American specimens as well. Doctor Hobart M.
Smith obtained data and photographs of certain Mexican specimens
in the Museum of Comparative Zoology at Harvard; Mrs. Helen T.
Gaige has furnished information on a specimen in the Michigan col-
lection. Dr. Emmet R. Dunn has placed certain data in my hands
and has offered valued criticism. I offer my gratitude to these per-
sons.

Genus Leptodeira Fitzinger

1826. Sihon (part.) Fitzinaer, N. Class. Rept., pp. 29, 31; Cope. Proc. Ac. Phila.. 1860,
266 (septei(trioitalis).

1837. Coronella (part.) Sclilegel, Phys. Serp.. H, 1837, p. .'JO (nifegcens).

1834. Leptodeira Fitzinger, Systeiiia Reptilium, j). 27. Type, "Dipsas aiiniilata Sclilegel"
[= Leptodeira rhombifera and Leptodeira aniiulata since this species of Schlegel is a composite
of Leptodeira rhombifera Giinther anrl Leptodeira annulata (Linne).

1861. Mei/alopa Hallowell (non Lacepetle), Pnic. Acad. Nat. Sci. Phila., 12, 1861, p.
488. Type, Megalops maculata.

1866. Leptodira Cope, Proc. Acad. Nat. Sci. Phila.. y. 127 (euitndation).

I have hesitated to accept Dunn's proposal to unite, with this
genus. Cope's Hypsiglena (type, ochrorhyiichd) . In his preliminary
paper he gives but little data for such a change. There are present
smaller series of teeth in the jaws; the fangs lack grooves; the elon-
gation of the snout anterior to the moutli is much greater than ob-
tains in typical Leptodeira; the tail is proportionally shorter, and
the scales differ in having only a single apical pit instead of paired
pits. Moreover, this genus has a distribution north of the Isthmus
of Panama, nearly coextensive with Leptodeira. I believe this genus
is a natural group worthy of generic recognition.

On the other hand Hypsiglena latifaseiata Giinther (which in-
cludes Leptodeira guilleni Boulengcr* as an absolute synonym) and
Hypsiglena discolor Giintlier lun'e jjaircd a))ical pits, shorter snouts,
witli fewer teeth that approach those of typical Leptodeira save that
the large back fangs are lacking in grooves. These forms are ex-
tremely rare, H. discolor being known only from the two cotypes,
and H. latifaseiata from only five specimens.

I regard these two forms as members of neither Hypsiglena nor
Leptodeira. I tlierefore propose for them a new generic designation.

* This form lacks grooved teeth, according to H. W. Parker, who at my request examined
the types and furnished data on all three mentioned forms. He concurs with Dunn's sug-
gestion that //. latifaseiata and L. guilleni are synonyms.

Taylor: Notes on the Genus Leptodeira 319

Le-ptodcira puticfatn (Peters)

(Phite XXX, fi«. 1)

1866. Crotaphopeltis punctatus Peters, Mon. Ber. Akad. Wiss. Berlin, 1860, p. 93 (type
description; type locality, "South Africa" [probably Western Mexico]).

1809. Leptodeira pacifica Cope, Pror. Acad. Nat. Sci. Phila., 1868 (1869), p. 310 (type
description; type locality, Mazatlan. Bishoff, Coll.); Giinther, Biologia Centrali-Americana,
Reptilia and Batrachia, 1895, p. 169; Boulenger, Cat. Snakes Brit. Mus., Ill, 1890, p. 19
(Presidio, near Mazatlan); Dimn, Proc. Nat. Acad. Sci., 22, 1936, pp. 691-094.

1887. Sibon pacificum Cope, Bull. U. S. Nat. Mus., No. 32, 1887, p. 67; and Proc.
U. S. Nat. Mus., XIV, 1892, p. 678.

1887. Leptodeira punctata Boulenger. The Zoiil., 1887, p. 178 (Africa).

1939. Leptodeira annulata punctata, .^niaral. Mem. Inst. Butantan, 1\'. \k 929, p. 204
(places Leptodeira nigrofasciata as a synonym).

1937. Leptodeira punctata Taylor, Univ. Kansas Sci. Bull. XXIV, 1936 (1937), pp. 526-
527.

A single male specimen of this rare snake was captured late at
night near a small railway bridge about a mile east of Mazatlan,
Sinaloa. The specimen was crawling along the bank of a small
rivulet which held water from a rain of the previous night.

It presents the following characters: Portion of rostral visible
above very narrow; frontal longer than its distance from the end of
the snout, shorter than the jiarietals; nostril very large, pierced
chiefly in the anterior part of the divided nasal; loreal small, as
high as wide; two preoculars, the upper very high, the lower minute;
two postoculars. both in contact with the single large anterior tem-
poral; posterior temporals two; diameter of eye equal to its distance
from the middle of the nostril. Upper labials 7-7, the sixth ex-
tremely large, the third and fourth entering the eye; anterior chin-
shields slightly wider, but no longer than the posterior; latter scales
separated from the first widened ventral by two pairs of small scales
and two single enlarged scales; lower labials 9-9, the first four touch-
ing the chinshields. Ventrals, 149; anal divided (preceded by a very
small median scale); suhcaudals, 70. Length, 516 mm.; tail, 130
mm.; Iietul width, 13 mm.; head length to angle of jaw, 19 mm.

Color in life. Above slightly reddish-brown with a series of black
spots extending to the tail on either side of the median line; and on
the side, one or two indefinite rows of irregular black fiecks tending
to form angular reticulations. Head brown; four small dark spots
on the posterior head scales; a small median black spot borders the
parietals and on either side of the nape are two large black spots
narrowly separated by' a yellowish area; no black bar behind eye;
labials very light tan; ventral surface cream.

Dunn's use of L. pacifica Cope for this form rather than L. punc-
tatus Peters is prompted by no new evidence.

320

The University Science Bulletin

Leptodeira splendida Giinther

(Plate XXX, fig. 2; Text fig. 1)

1895. Leptodeira splendida Giinther, Biologia Centrali-Aiiiericana, Reptilia and Batrachia,
May, 1895, jip. 168, 170, pi. LIII, fig. B (type description; type locality, Izucar, Puebla,
Me.xico).

1896. Leptodira pcmoiiata (part.) Boulenger. Cat. Snakes British Mus., Ill, 189f), p. 9.3
(types of splendida): Mocquard (part.), Mission Scientifique au Mexique, livr. Id, 1908, pp.
903-904.

1936. Leptodeira rhonibifera (part.) Dunn, Proc. Nat. .\cad. Sci., vol. 22, 1936, pp. 691-
693.

Four specimens of this species are in the collection. Nos. 5177-
5179, from a point 12 miles south of Puente de Ixtla, Morelos (E. H.
Taylor, collector), and No. 5478, from Cuernavaca, Morelos (H. M.
Smith, collector).

Fig. 1. Li ptodeira .splendida (^iiinther, EHT-HMS, No. 5179, twelve miles south
of Puente de Ixtla, Morelo.'^, Mexico (enlarj>ed) .

This species has been confused with lx)tli personata and rhombi-
fera. It occurs in Morelos, and Puebla. It may be characterized
by the presence of a stripe or bar on the nape; tlie peculiar head
pattern; the presence of three preoculars (likewise typical of septen-
trionalii^, frenata and polysficta) ; a reduced number of small dorsal
spots, 19-25; 21 scales about middle of the body; ventrals, 165-168;
subcaudals, 76-80. The posterior fangs are deeply grooved. The
spots reach on the sides to the third or fourth scale rows, often tend-
ing to break medially or to become confluent. The following table
shows the variation obtaining in this form.

Taylor: Notes on the Genus Leptodeira

321

Table of data and measurements in mm. of Leptodeira splendida Giinther

Number

5178

5179

5478

5177

Type

Sex

cf

d'

?

cf

9 ?

168
80

167
79

167
76

166
79

165-66

Subcaudals

76

3-3
2-2

3-3
2-2

3 3
2-2

3-3

2-2

3-3

Postoculars

2-2

1+2+3

1+2+3

1+2 43

1+2+3

1 +2 +3

Soalp formula

21-21-17
490

19-21-16
473

21-21-17
362

21-21-16
273

—21 —

Length, total

650

129

122

89

70

162

Spots, body

25

23

19

23

20

Spots, tail

20 +

20 +

15 +

14 + +

Giinther's type specimen is a relatively large one and the pattern
on the dorsal surface of the head has more or less disappeared. (It
is less distinct in my No. 5178.) One of the most striking differ-
ences between this form and rhombijera is the presence in males of
keels on the scales along the posterior fourth or fifth of the body and
on the base of the tail. In females the keels are barely discernible,
which probably accounts for the fact that they are not recorded in
the type. The constancy of three preoculars is likewise a pertinent
character. The grayish or grayish-brown spotting is likewise in
contrast to the brown or blackish-brown coloration of typical rhom-
bijera.

The intercalated lateral spots are present but small, often some-
what elongate. The body is not compressed; the chinshields are of
about equal length. The preocular touches the frontal in two cases;
they are separated in two cases.

I believe this form to be more closely related to Leptodeira bres-
soni, described herein, than to rJiombijera.

Leptodeira bressoni sp. nov.

(Plate XXXI, fig. 4; Text fig. 2; Plate XXXIII, fig. 4)

Type. EHT-HMS, No. 5172, collected at Hda. El Sabino, about
20 miles south of Uruapan, Michoacan. Don Julio Raymond Bres-
son, collector.

Paratypes. Nos. 4617, El Sabino, Michoacan, July 21-28, 1935,
H. M. Smith, collector; 4619, near Queseria, Colima, June 18, 1935,

21—4141

322

The University Science Bulletin

H. M. Smitli, collector; 5173, Hda. El Sabino, 1935, Don Julio Ray-
mond Bresson, collector.

Diagnosis. Related to L. splendida, but differing from it in color
and scale characters. Scales in 19-21 rows; a dark nape stripe; the
lines behind the eyes tending to join the first nuchal dark mark and
usually separated by only a very short distance.

Ventrals, 168 to 182; subcaudals, 81 to 90; upper labials, 8-8; lower
labials, 10-10; temporals, l+2-|-3; dorsal scales keeled on pos-

,*3^r-v.

Fig. 2. Leplodeira bresso7ii sp. nov. Paratype. EHT-HMS, No. 5173. Hda.
El Sabino. Uniapan, Michoacan (enlarged).

terior fourth or fifth of body; a bar on the nape of the neck, but not
joining the first dorsal mark. Dorsal spots, 32-38; lateral inter-
calated spots prominent.

Description of the type. Rostral barely visible from above, its
width once and one third its height; internasals typically small, their
areas about "one third that of prefrontals; latter large, distinctly
wider than long; frontal much wider than long, its length equal to
its distance from tip of snout; parietals typical, their length about
one and one half times their width; nasal divided, or at least par-
tially so; loreal slightly longer than high; three preoculars, the upper
more than double size of middle one, not touching frontal; lower
preocular (subocular) very small, separating third labial from eye;

Taylor: Notes on the Genus Leptodeira

323

supraocular widened posteriorly; temporals, 1+2+3; two postoc-
ulars; upper labials, 8-8, fourth and fifth entering orbit, having the
following order of size, 1, 3, 2, 4, 5, 8, 6, 7; lower labials, 8-9 (ab-
normal, due to fusion of the ninth and tenth on one side) ; the pos-
terior chinshields are longer than the anterior, pointed behind, sep-
arated from first widened ventral by two pairs of scales. Scale for-
mula, 19-21-17; scales of the dorsal rows with keels on posterior
fourth of body, growing more pronounced posteriorly above the base
of tail; some of the keels are not continuous; some have a slight,
knoblike termination. Ventrals, 171; subcaudals, 84; anal divided,
lengthened abnormally, preceded by an unequal pair of scales.

Color. The general ground color is a rosy flesh. The body has a
series of spots continuous across the middle of the back or broken
medially and tending to alternate, but usually in contact. When
broken the parts are quadrangular. The number of spots and pairs
of spots on body, 38; on tail, 24+. On the intervening areas of
ground color, there is an indistinct darker band which traverses the
body from the first or second scale rows. The primary dark blotches
reach to fifth scale row, and below these are indistinct blotches
reaching to the ventrals. These intercalated markings are visible in
the very young, the lower parts being darkest. First dorsal blotch
forms a V-shaped mark, which is about three scales wide on the
middorsal line; the bar on the nape beginning at the parietals ex-
tends back between the anterior arms, but does not touch the spot;
a black bar from the eye runs to posterior edge of the eighth labial;
this is followed by a small dark spot at a short interval. The top

Table of scale data and measurements in mm. of Leptodeira bressoni

Number

Sex or age

Ventrals

Subcaudals

Upper labials

Lower labials

Preoculars

Postoculars

Temporals

Scale formula

Length, total (mm.)
Length, tail (mm.) .

5172

&

171

84

8-8

8-9

.3-3

2-2

1+2+3

19-21-17

481

124

5173

168

90

8-8

10-10

3-3

2-2

1+2+3

19-19-17

430

118

4617

177

81

8-8

10-10

3-3

2-2

1+2+3

21-19-17

.335

89

4619

yg.

169

82

8-8

10-10

3-3

2-2

1 +2 +3

21-21-17

210

53

MCZ

11411

9

172

71

8-8

10-10

3-3

21-19-17
372
107.5

USNM
46459

9

182
78
8-8
10-10
3-3
2-2

21-21-17
548
128

324 The University Science Bulletin

of the head is variously mottled with black, but is injured so that
no definite pattern can be discerned. The ventral surface is rosy
flesh; the lateral edges of the ventrals with scattered dark pigmenta-
tion; the anterior chin scales likewise with some dark pigment;
subcaudals with A^ery slight pigmentation near tip of tail.

Variation. Dr. Emmet R. Dunn called my attention to the fact
that MCZ, No. 11411, Colima, Mex. (plate XXXIII, fig. 4), proba-
bly belonged to this form. An examination proves that such is the
case. It agrees very well, falling within the variation observed in the
Michoacan specimens. Being a female, the ventral count is some-
what lower. The spotting on the body is irregular (31 on one side,
34 on the other, due to the fact that certain spots have broken apart ).
The tail has 12 spots on one side, 13 on the other. Dorsal scales are
keeled on posterior part of body. Stippling present on chin and on
under side of tail.

I am likewise associating with this species USNM No. 46459,
Plomosas, Sinaloa; Nelson and Goldman, collectors. It is a female,
with a slightly reduced subcaudal count ; the general head and neck
pattern is generally similar but somewhat obscured, due to injury to
the top of the head. The most pertinent difference is that the spots
are transverse, larger, wider, not tending to break medially. They
extend to the second scale row, with some smaller spots on the first
row below the lateral ends of tlie dorsal spots. The large spots are
two and one half to three scales long on dorsal surface. The spots
are brownish-gray, darker on their edges.

The number of spots on the tail is 18-19, fewer than in the type.
The scales are keeled on the posterior fourth of the body. The fangs
are grooved. The frontal is shorter than its distance to the tip of
the snout.

It is probable that with a considerable series from this region it
might be possible to separate this from the typical L. bressoni as a
subspecies.

Remarks. This form has the dorsal spots varying in number be-
tween 32 and 38. In the youngest specimen. No. 4619, the flesh
color is less pronounced.

This species is named for Don Julio Raymond Bresson, Dr.
Smith's host at Hda. El Sabino, who showed him every courtesy and
contributed much in the way of assistance in collecting, and pre-
sented him with many specimens.

The species differs from Leptodeira splendida in having a different
pattern on the head, larger number of spots on the body, 32-38

Taylor: Notes on the Genus Leptodeira 325

(19-25 in L. splendida) , the presence of a heavy reticulation on the
ground color, and a somewhat higher ventral count, 167 to 182, in-
stead of 165 to 168.

Leptodeira mystacina Cope

18G9. Leptodeira mystacina Cope, Proc. Anier. Philos. Soc, XI, 18G9, 151 (type descrip-
tion: type locality, "Western Region of Mexico near the isthmus of Tehuantepec." Two speci-
mens, F. Sumichrast, collector); GUnther, Biologia Centrali-Americana, Rept. Batr., May,
1896. p. 1G9 (description of specimen not the type); Dunn, Proc. Nat. Acad. Sci., 22, No.
12, Dec, 1936 (Acapulco, Guerrero, and Tapanatepec, Oaxaca).

1887. Sibon mystacinum Cope, Bull. U. S. Nat. Mus., No. 32, 1887, p. 67 (Nicaragua and
West Tehuantepec).

1892. Sibon nigrofasctattun Cope, Proc. U. S. Nat. Mus., XIV, 1892, p. 678 (Unites L.
nigrofasciata and L. vrystacinum and gives generic key); and Ann. Rop. U. S. Nut. Mus.,
1898 (1900), p. 1107.

1896. Leptodeira nigrofasciata (part.) Boulenger, Cat. Snakes British Mus., vol. Ill, 1896,
p. 92 (specimen from Tehuantepec) ; Mocquard, Mission Scientifique au Mexique et dans
TAmerique Central, livr. 16, 1908, pp. 900, 901 (description of a Tehuantepec specimen).

Opinions as to the distinctness of this form have differed. Cope,
1892, regards his own species a synonym of L. nigrofasciata Giin-
ther, which was described some months earlier than L. mystacina
Cope. Giinther, who had available a specimen from Tapana, Te-
huantepec, likewise collected by Sumichrast, regarded the species
distinct from his L. nigrofasciata, in spite of Cope's action, which
Giinther mentions in a footnote. Moccpard (1908) with three speci-
mens collected by Sumichrast in Tehuantepec, either on his own
judgment or on Cope's action, regards them as L. nigrofasciata.
Dunn, 1936, with 13 Nicaraguan, and 2 Costa Rican specimens, to-
gether with Cope's types of L. mystacina and three additional speci-
mens from Guerrero and Oaxaca, Mexico, accepts them as distinct
species. The following distinguishing characters are presented in the
key:

L. nigrofascinta : 15-18 bands; ventrals, 168-172; Nicaragua and Costa Rica.
L. mystacina: 10-12 bands; ventrals, 187-196; Guerrero and Oaxaca.

The higher ventral count of L. mystacina as well as the reduced
number of bands seem to warrant the retention of L. mystacina.
Certainty as to whether intergradation occurs will have to await
more extensive collecting in territory between the known ranges.

Tlie following data were taken from specimens in the U. S. Na-
tional Museum.

The cotypes (Nos. 30339-40, Tehuantepec, Mexico; Sumichrast,
collector) present the following characters:

No. 30339: Rostral visible above as a line; frontal narrow, elon-
gate, distinctly longer than its distance from end of snout; parietals
much widened anteriorly; prefrontals very large, forming an angle

326 The University Science Bulletin

laterally on right side; preoculars 2, 2, the upper very large — much
widened and much elongated, forming a suture with frontal, more
or less angular on its anterior edge, pushing between prefrontal and
loreals; upper labials, 8 (right), 7 (left, 2 anterior labials abnormally
fused) ; the sixth labial on right side touches upper postocular, but is
minutely separated from the parietal; the fifth labial touches the
parietal on the left side; lower labials, 10-10; postoculars, 1, 2;
temporals 1+1+2+3; scale formula, 23, 19, 19, 17; ventrals, 187;
caudals, 69; 11^/2 spots on body counted from one side, 13yo on the
other; posterior chinshields separated; the head does not havo the
scales outlined with lighter color, and there is a very slight segrega-
tion of pigment at the anterior edges of the parietals and toward
their posterior parts; a dark line in front of eye, continued behind
eye to angle of mouth, and a second line below eye, diagonally to
labial border, separated from the other dark line by a light line;
upper labials with dim markings; on body there is a segregation of
pigment to the edges of the scales of the dark spots.

No. 30340: General characters the same; preoculars. 2-2; post-
oculars, 2-2; the labials narrowly separated from tlie parietals;
temporals, 1+2+3; second chinshields partially separated; IIV2
spots on body; 4:'^^ on tail, no evidence of pigment segregation on
dorsal surface of head. Ventrals, 194; subcaudals, 65.

No. 46551: (Nelson and Goldman, Acapulco, Guerrero, 1895).
Agrees with the preceding specimens in general characters. Scale
formula, 23, 17, 19, 17; preoculars, 2, 2; postoculars, 1, 1, the lower
apparently fused to the fifth labial, which rises as high as middle of
the eye; temporals, 1+2+3; upper labials, 8-8; lower labials, 9-9;
chinshields of equal lengih, the second pair separated; ventrals, 192;
subcaudals, 65. Body with 11 deep lavender spots, the two sides not
the same, since one half of each spot has moved forward nearly a
half its length; each scale of the dorsal spots has a lighter center,
thus giving the marking a reticulated appearance; each spot has a
darker border; intervening ground color distinctly lighter, lightest
where it contacts the dark spots; three spots on tail, not strongly
distinct from intervening spaces. A brown line from nostril to eye,
and from eye to jaw angle; a light line along the upper part of upper
labials anteriorly, which turns down across posterior labials; a
strongly defined dark line from below eye on fifth and sixth labials;
chin powdered with dark pigment which forms indistinct spots; the
lateral coloration encroaches on the ventrals ; ventral surface of tail
pigmented. Top of head stippled lightly with lavender-brown.

Taylor: Notes on the Genus Leptodeira 327

Leptodeira dunckeri (Werner)

1913. Leptodeira dunckeri Werner, Mitt. Nat. Mus. Hamburg, 1!»13, 30, pp. 28, 29 (type
description, type locality, "Mexico oder, Venezuela"; also a key to the American species of the
genus).

1929. Leptodeira avmilata anmilnta, (part.) Amaral. Mem. Inst. Butantan, 4. 1929, p. 204.

1930. Leptodeira septentrionalis macuiata (part.) Dunn, Proc. Nat. Acad. Sci., 22, 1936,
p. 091.

Dunn has regarded this species as being probably Mexican and
has placed it as a synonym of Leptodeira septentrionalis macuiata
(Hallowell) .

The form is diagnosed as having twenty-one scale rows, 183 ven-
trals, seven supralabials, one preocular, one postocular; whitish with
twenty-five dark red-brown transverse bands which reach to the
ventrals, narrowing on the sides, some of which bands are in contact
dorsally, tending to form a zig-zag band. Between the dark bands
near the ventrals are larger dark flecks. Two spots, side by side,
on the parietals; snout darkly punctate: upper labials flecked with
dark color; underside white.

It may be noted that the form differs from Mexican Leptodeira
macuiata in the presence of a single postocular instead of two, one
preocular instead of two (one occasionally in western "personata") ,
seven instead of eight upper labials, 21 instead of 23 scale rows
around the body (rarely 21 in macuiata). The ventral count is
higher than any Mexican specimen of macuiata (Dunn gives 186 as
maximum. I have not seen this specimen) .

I feel that the evidence available does not wholly justify associa-
tion of this name to the synonymy of Leptodeira macuiata regardless
of whether the specimen originated in Venezuela or Mexico. The
very brief description suggests a specimen more closely related to
L. yucatanensis malleisi than to L. macuiata. It has the scale rows
of this form; one postocular (occasional in malleisi), the same num-
ber of bands, a ventral count within the range of the latter, and
paired spots on the parietals, usually present in malleisi. However,
this cannot be satisfactorily settled until the type of L. dunckeri is
studied in greater detail.

Leptodeira yucatanensis malleisi Dunn and Stuart

(Te.xt Fig. 3)

193.5. Leptodeira yucatanetisit malleisi Dunn and Stuart, Ofc. papers, Mils. ZooL, U. of
Michigan, No. 313, May 29, 1935, pp. 1-4 (type description; type locality, Tuxpcna, Cam-
pech', Mexico); Stuart, Univ. Michigan, Mus. Zool., mis. publ. No. 29, Oct. 1. 1935, pp. 8,
■24, .">3 ; Claire, Carnegie Inst. Washington publ. No. 457, p. 302; Dunn, Proc. Nat. Acad. Sci.,
22, 193G, pp. 691, 696.

Two specimens of this form collected by Dr. Hobart Smith are
in tlie collection, Nos. 11618 j from Encarnacion, Campeche, and

328

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11619 c^ , from Pital, Campeche, collected on October 14, and Oc-
tober Ki, 1936. These have the following characteristics respec-
tively: Scale formula. 19-21-15, 19-21-15; ventrals, 179, 188; siib-
caiidals, 79, 67; supralabials, 8, 8; lower labials, 10-10; preociilars,
2-1, 2-2; postoculars, 2-2, 2-2; temporals, 1+2+3, 1+2+3; total
length, 431 mm., 627 mm.; tail, 97 mm., 138 mm.; spots on body,
33, 28.

Fig. 3. Leptodeira yucatanens'is nialleisi Dunn and Stuart. EHT-HMS,
No. 11619 9. Pital, Campeche, Mexico (enlarged).

In No. 11619 the ground color and ventral surfaces are quite red-
dish. The black blotches are lighter in their centers and bordered
by a thin whitish line. Many of the blotches are united, forming a
zig-zag line on back. The blotches reach to the second or third
scale row. Intercalated spots low on the side are distinct. There
are also spots below the ends of the blotches; a nape stripe, not
reaching the first nuchal blotch, and a pair of spots on the parietals.
The lines from the eye reach the first nuchal blotch.

The smaller specimen is lighter pink on the ground color and
ventral surfaces, and the spots are regular; the paired spots on the
parietals are surrounded by a light ring.

Taylor: Notes on the Genus Leptodeira 329

Leptodeira yucatancnsis yucatancmis (Cope)

Leptodeira annulata var. Cope, Pioc. Acad. Nat. Sci., Philadelphia, 186R, p. 127.

Sibon annulata yucatanensis Cope, Bull. U. S. Nat. Mus., 32, 1887, p. 67 (type de-
scription by reference to Cope, 1866; type locality, Yucatan).

Leptodeira yucatanensis yucatanensis Gaige, Carnegie Inst. Washington Pub. No. 4;)7,
Feb. 5, 1936, p. 302 (Chichen Itza).

I have not examined specimens of this form.

Leptodeira septcntrioncdis (Kennicuttj

(Plate XXXr, fig. 3; Te.xt fig. 4)

Dipsas septentrionahs Kennicott, in Baird, Report Mexican Boundary Survey, 2, Rept.,
1859, p. 16, pi. 8, fig. 11 (type description, type locality, Matamoras, Tamaulipas, Mexico,
and Brownsville, Texas; collected by Lieut. Couch and \'an \'liet); Cope, Proc. Acad. Nat.
Sci. Phila., 1860, p. 266.

Sibon annulatum septentrionale Cope, Check-list Batr. and Rept. N. Amer., 1S7.5, p. 38;
Bull. U. S. Nat. Mus., 1887, p. 67 {part.).

Sibon septentrionale (part.) Cope, Proc. U. S. Nat. Mus., XIV, 1891, iip. 677-678 (Key;
ranges from Panama to Cameron county, Texas); Ann. Rep. U. S. Nat. Mus., 1898 (1900),
pp. 1007-1008, fig. 316 (Matamoras, Mex.).

Leptodeira septentrionahs Stejneger, Proc. U. S. Nat. Mus., XIV, 1891, p. 50.5.

Leptodeira annulata (part.) Giinther, Biologia Centrali-Americana, Reptilia, 1895, p. 170.

Leptodeira septentrionalis Boulenger, Cat. Snakes British Mus., Ill, 1896, p. 93.

Sibon septentrionalis Strecker, Baylor Bull.. XVIII. No. 4, Aug. 191. J, p. 41 (Cameron
county, Texas).

Leptodeira septentrionalis septentrionalis Dunn., Proc. Nat. Acad. Sci., vol. 22, 1936, pp.
692. 697 (Cameron county, Texas, to Tampico, San Luis Potosi. Mexico, Zacualtipan, Hidalgo,
Mexico).

This Large, robust form is represented in our collection by three
specimens: No. 4615, seven miles west of Victoria, Tamaulipas;
4616, Hda. La Clementina, Tamaulipas; and 4652, uncertain locality.

This form may be diagnosed by the presence of three preoculars
(pre- and suboculars) ; by the large black blotches touching or
reaching near the first scale row-; the high number of ventrals and
subcaudals (186 to 197; 63 to 79, combination of data given by
Dunn [1936] and my own data), and the presence of more or less
pigment on the posterior edges of the ventrals; the scale count re-
duces to 15 in front of the anus. There is a black spot on the pos-
terior edges of the parietals, usually confluent with an indistinct
head pattern.

Data on the three specimens, Nos. 4615, 4616, and 4652, respec-
tively, follows: Sex, ^ , ^ , ^ ; ventrals, 193, 191, 186; subcaudals,
79, 81, 77; supralabials, 8-8, 8-8, 8-8; lower labials, 10-10, 10-10,
10-10; preoculars, 3-3, 3-3, 3-3; postoculars, 2-2, 2-2, 2-2; temporals,
1+2+3, 1+2+3, 1+2+3; scale formula, 26-21-23-15, 25-21-23-15,
26-21-23-15; rhombs on body, 23, 23, 24; spots on tail, 14, 12, 15;
total length in mm., 785, 360, 256; tail, 180, 83, 55.

The black of the large spots is rather dense and does not show

3:30

The University Science Bulletin

Fig. 4. Leptodcira septentrionalis (Kennicott). EHT-HMS, No. 4616.
La Clementina, near Forlon, Tamaulipas, Mexico (enlarged).

Hda.

flecks of white. The posterior teeth of the maxillae are strongly
grooved.

Doctor Dunn has very kindly supplied data on specimens of L.
septentrionalis. He lists from Zacualtipan, Hidalgo, ANSP Nos.
11662 5 , 11663 ^ , 14775 ^ . They have the following ventral counts,
respectively: 197, 196, 197. All have three preoculars. His speci-
men of maciilata from this localit}^, ANSP No. 14744 ^ , has only
167 ventrals and two preoculars.

A series of L. septentrionalis from Tuxpan, Veracruz, in the U. S.
National Museum, Nos. 25206 $ , 25207, 25209, 25210, 25211 (the
last four young) , have the following ventral and subcaudal counts,
respectively: 192, 80; 193, — ; 191, 82; 201, 73; 201, 73. All reduce
the scale count to 15 in front of anus. The spots on the body vary
between 27 and 33. The typical head marking and the small
postparictal black spot are present. The body spots are less scjuare
on the sides, tending to narrow more than in more northern forms,
and the interspaces are somewhat wider. All have three preoculars.

Two of the cotypes (USNM No. 4267, two specimens, Matamoras,
Tamaulipas, Mexico) are small. Each has three preoculars. The
ventral counts are 186 and 189; the subcaudal counts, 77 and 64.
Scale formulae, 17-21-15; 21-23-15. Another cotype mentioned,
No. 2288, Brownsville, Tex., is not to be found. It may be the speci-

Taylor: Notes on the Genus Leptodeira 331

men figured by Kennicott {loc. cit.). This specimen is abnormal in
having several undivided subcaudals following the anus.

USNM No. 4273, from Matamoras, Tamaulipas, is the only speci-
men examined having the scale count of 17 one-half inch in front of
vent. In all others it is reduced to 15.

Dunn (1936) has proposed the placing of Leptodeira septentriona-
lis as a subspecies of L. maculata, and suggests that L. maculata re-
places the form off the plateau. The data here presented show a
much greater overlap of the ranges than was hitherto suspected and
in this range there is no evidence that the characters of the one- ap-
proaches the general characteristics of the other. Because of this
wide difference in the count of ventral scales, the presence of three
instead of two preoculars, a different head pattern, and the pigmenta-
tion on the ventral scales, I regard the two forms specifically distinct.

Despite the fact that Leptodeira polysticta shows certain char-
acteristics in common with L. septentrionalis, i. e., scale counts within
the same ranges, three preoculars and pigmentation on the posterior
edges of the ventral scales, I am thoroughly convinced that the two
forms are only remotely related.

Leptodeira annulata polysticta Giinther

(Plate XXX, fig. 3)

Leptodeira annulata (^part.) Giinther, Cat. Col. Snakes British Mus., 1858, p. 160.

Leptodeira polysticta Giinther, Biologia Cent rali- Americana. Reptilia and Batrachia, May,
1895, p. 172, pi. LV, fig. a (type description; type locality, Mexico, Jalapa, Oaxaca, Yucatan;
British Honduras, Belize; Honduras, and Panama).

Leptodeira albofusca {part.) Boulenger, Cat. Snakes, British Mus., 1896, pp. 95-97;
Mocquard (.part.), Mission Scientifique aui Mexique, etc., Rept., liv. 16, 1908, pp. 905-906;
? Werner Abh. Akad. Wiss., Bd. XVII, Abt. II, p. 348 (Coban, Guatemala); ? Shattuck,
et al., The Peninsula of Yucatan, 1933, p. 575 (Yucatan).

? Leptodeira annulata Dunn and Emlen, Proc. Acad. Nat. Sci. Phila., LXXXIV, 1932, p.
32 (Rancho El Jarrae, Honduras); Sumichrast, Arch. Sci. Phys. Natur., 46, 1873, p. 246.

Leptodeira annulata polysticta Stuart, Misc. Publ. No. 29, U. of Michigan, Mus. Zool.,
Oct. 1, 1935 (La Libertad, Guatemala); Gaige, H. T., Carnegie Inst. Washington, Publ. No.
457, Feb. 5, 1936, p. 302 (Chichen Itza, Yucatan, Mexico); Dunn, Proc. Nat. Acad. Sci., 22,
1926, pp. 691, 693 (Colima, Southern Veracruz, Yucatan, to Peten, Guatemala and Eastern
Honduras).

This subspecies is represented in the collection by the following
specimens: No. 4618, Acultzingo, Veracruz, elevation 6,000 feet, E.
H. Taylor, collector, 1932; 4620, Hda. Paso del Rio, Colima, H. M.
Smith, collector, 1936; 11616, Pital, Campeche, and 11617, En-
carnacion, Campeche, H. M. Smith, collector, 1936. I have also
examined a specimen in the collection of the University of Kansas,
No. 8486, from an unknown locality (collected in Pocatello, Idaho,
by Wayne AVhitlow from a bunch of bananas), and a series of
specimens in the United States National Museum.

This elongate, slender species is characterized by the large number

332

The University Science Bulletin

of small dorsal blotches on the body with a more or less distinct
series of small lateral spots alternating with the dorsal spots ; a black
bar on the nape not connecting with the first dorsal blotch; a dim,
dark bar behind eye, not connecting with the first dorsal bar; three
preoculars (preoculars and suboculars) ; the high ventral count is
characteristic of the species (198-211 ventrals; 80 to 102 subcaudals
[according to Dunn, 1936, 193-211, 75-102]. I suspect the lower
counts are from U. S. N. M. 25206-7, 25209-11, from Tuxpan, which
I refer to L. septentrionalis) .

The body is more or less compressed in all the specimens, but in
the Colima specimen it is especially compressed, leaving a sharp
dorsal ridge evident. This does not appear to be due to emaciation.
This character is absent in the other specimens. In all there is some
distribution of fine pigment on the ventrals, especially on the
posterior half of the body. It is more dense under the tail. The top
of the head is pigmented more or less uniformly, but there is no
definite pattern.

The ground coloration of the Veracruz and Campeche specimens
is a light brown, sometimes flesh or pinkish brown; ventral surface
flesh color. The Colima specimen is gray, the ventral coloration
being nearly white. The first dark nuchal blotch is V-shaped or U-
shaped, usually not more than two or three scales wide on dorsal line.

The following table gives variational data, presented by the speci-
mens:

Table of data for Lcptodeira annulatn polysticla Giinther

Number

4620

4618

11617

11616

8486

Sex or age

Scale formula

Ventrals

2o-21-£S-l7

209

80

8-8

10-10

3-3

2-2

1+2+3

845

186

'60

29

yg.

26-23-;2.i-17

198

84

8-8

10-11

3-3

2-2

1 +2+3
1+2+4

340

76

45

22

yg.

26-21-^5-17

208

86

8-8

10-10

3-3

2-2

1+2+3

347
78
52
24

yg.

26-21-J.J-17

205

83

8-8

10-11

3-3

2-2

1+2+3

437
97
57
23

9
26-21-J.5-17
203

Subcaudals

Upper labials

Lower labials

Preoculars

Postoculars

Temporals

Total length fmm.)

Tail length fmm.)

Spots, body

Spots, tall

90

8-8

10-10

3-3

2-2

1+2+3

878

205

56

25

Taylor: Notes on the Genus Leptodeira 333

A series of specimens in the United States National Museum from
Veracruz (7088, 30207, 30208, Orizaba, Sumichrast, collector; and
65154, hills West of Veracruz, Sartorius, collector; 30508, Veracruz,
Sumichrast, collector) together with my specimen from Acultzingo,
Veracruz, differ from the western Mexican specimen from Colima
in having the dorsal spots number only 38 to 54 instead of 60. Two
of these have only 2 preoculars, the small lower preocular being ab-
sent. The specimens from British Honduras, USNM No. 26058,
Tela, Honduras, USNM 64683, and my specimens listed from
Campeche have the dorsal spots varying between 52 and 57.

Since I discern no significant differences in squamation or general
color characteristics in these specimens, I shall await more material
before considering the possibility of separating the eastern and
western Mexican forms.

Leptodeira frenata (Cope)

Sibon frenatum Cope in Ferrari -Perez, Proc. U. S. Nat. Mus., 9, 1886 (1887), p. 184
(type description, type locality, Jalapa, Mexico).

Leptodeira annulata ■personata (part.), Aniaral, Mem. Inst. Butantan. IV, 1929, p. 204.
Leptodeira frenata Dunn, Proc. Nat. Acad. Sci., 22, 1936, pp. 692, 696.

This form has not been rediscovered, and the type is lost. The
original description follows:

"Scales in twenty-three longitudinal series. Body rather slender, tail rather
short, head very di.stinct and depressed. Superior labials nine, eye resting on
the fourth and fifth, and onlj' separated from the third by the small inferior
preocular. All are higher than long, excepting the eighth and ninth, which are
longer than high; the sixth and seventh are the largest. Inferior labials, eleven.
Postgeneials much longer than the pregeneials. Loreal plate subquadrate;
oculars, 2-2; the superior anterior not reaching the frontal plate. Temporals,
1 + 2 + 3. Frontal twice as long as wide, with parallel sides. Occipitals
moderate, reaching to above middle of eighth superior labials. Gastrosteges,
188; anals, 1-1; urosteges, 69.

"Colors: Above black, below white. At distance of from si.x to nine scales,
narrow cross-bands of one scale in width rise from the abdominal border color,
and meet or terminate in alternating positions on or near the middle line of
the back. These bands are more or less gray, sometimes darker in the middle.
The top of the head is gray, densely mottled with blackish, leaving a crescentic
space of light gray between a black spot behind the head-shields and the be-
ginning of the black of the superior surfaces. A broad, black band passes
downwards and posteriorly from the eye, and cro.ssing the angle of the mouth,
covers the side of the neck and unites with the black of the following regions.
The superior labials are light gray, with black borders. The dark borders of
the inferior labials are less distinct.

"Total length, 30.5 mm.; of tail, 66 mm.; of head to canthus oris, 11 ram.
No. 298 [of the Comi.^ion Geografica Exploradora de Mexico collection ex-
hibited at the New Orleans Exposition] Jalapa, Mexico.

334

The University Science Bulletin

"This species is nearest the .S. personatum Cope from Mazath'm, although
the coloration is very different. That species had but one preocular, eight
superior labials, etc." — E. D. Cope.

There is a possibility that the type of this species was returned to
Mexico and it may be rediscovered there. There is no evidence that
it was entered in any eastern museum. Cope studied the specimens
at New Orleans according to a statement {loc. cit.) of Ferrari-Perez
(p. 182).

Leptodeira smithi sp. nov.

(Plate XXXI, fig. 2)

Type. EHT-HMS No. 5187. Collected Hda. El Sabino, 19 miles
south of Uruapan, Michoacan, Mexico, August 2, 1936. Hobart
Muir Smith, collector.

Paratijpcs. EHT-HMS No. 4633, July 21-28, 1935; No. 5186,
July 21, 1936, and No. 5188, August 2, 1936, all collected by Hobart
M. Smith at Hda. El Sabino, Michoacan; FMNH No. 985, Balsas,
Guerrero, S. E. Meek, collector.

Diagnosis. Fangs grooved; body with 12-15 dark bands, at least
double the width of the intervening light spaces; no nuchal line;
body not, or but slightly compressed, lacking all trace of keels in

Fig. 5. Lcplodcvm .smithi sp. nov. Type EHT-HMS, No. 5187. Hda, E
Sabino, Uruapan, Michoacan, Mexico (enlarged).

Taylor: Notes on the Genus Leptodeira 335

males and females; 23 (or 21) scale rows; ventrals j , 167-168; ^"^ ,
169-173; caudals ^ , 17-79, 5 , 70; preoculars, 2; postoeulars, 2;
temporals, 1 + 2 + 3; upper labials, 8; lower labials, 10.

Descri'ption of the type. Head rather flattened, distinct from
neck; rostral not or scarcely visible above, 3.2 mm. wide by 2 mm.
high; internasals about as long as broad, their length along median
suture, 2 nun.; prefrontals slightly wider than long, 3 X 3.2 mm.;
frontal a little shorter than its distance from end of snout, widest
anteriorly, the sides not parallel; trace of a groove in the middle
line of frontal, with a small regular depression (roughly arrow
shaped) at the posterior end of the groove; parietals 6.4 mm. X 4.2
mm., touching only upper postocular; median scale, following median
parietal suture only slightly enlarged; nasal large, at least partially
divided, the nostril near the middle of the scale, the posterior moiety
larger than the anterior; loreal large, longer than high, the lower
border much longer than upper; upper preocular very large, irregular,
touching the frontal above and the third labial below; lower preocu-
lar (subocular) small, square, separating the third labial from the
eye; supraoculars wider posteriorly than anteriorly; a large anterior
temporal touching both postoeulars, followed by two, the upper the
larger; these followed by three temporals, the lower of the three
extending half its length behind the eighth labial; eight upper
labials; fourth and fifth form lower edge of orbit, arranged in the
following ascending order of size: 1, 3, 4, 2, 5, 8, 6, 7. The scale
following the last upper and lower labials is enlarged; mental small,
nearly an equilateral triangle; lower labials ten, five touching the
anterior chinshields; the latter scales equally as long and somewhat
larger than posterior; two pairs of scales between second chinshields
and the first ventral; five rows of scales between the last lower
labial and the third ventral; scale formula, 25, 19, 23, 17; ventrals,
169; anal divided; caudals, 70. Total length, 510 mm.; tail, 115
mm.; proportion of tail to body length, .22.

Color. Ground color dirty brownish, the lateral scales of the
ground color being edged with blackish, while those on the dorsal
part are clear light brown, almost lacking black pigment. Head
above blackish or blackish brown, with a trace of whitish bordering
most of the head scales, and slightly lighter areas on frontal and
parietals; temporal scales usually light-edged, with dark centers;
two or three labials with dim dark spots in center; upper edge of
posterior upper labials crossed by a dark bar from eye, which termi-
nates on the eighth labial; first light band on occiput about four and

336

The University Science Bulletin

one half scale-rows wide; first black band largest, 15 scale-lengths
wide; succeeding bands vary from a width of 10 scales to 8 scales;
the smaller bands are more posterior. Ventral surface immaculate,
save for a peppering of pigment on the anterior part of chin and a
few flecks under posterior part, of tail. The bands reach laterally
and cover part of the scales of the first row.

Table of Measurements in mm. and data of Leptodeira smithi sp. nov.

Number. . .

Sex

Ventrals. . . .
Subcaudals. .
Preoculars. . .
Postoculars . .
Temporals . . .
Scale formula
Length, total
Length, tail . .
Spots, body . .
Spots, tail . . .

4633

&

167

79

2-2

2-2

1+2+3

21-23-17

262

68

14

7

5186

a"

169

77

2-2

2-2

1+2+3

21-21-15

408

97

12

5187

9

169

70

2-2

2-2

1+2+3

(19-21)-23-17

510

115

14

7

5188

9
173
?

2-2
2-2

1+2+3

20-23-17

434

22 (broken)

15

7

Variation. Outside of variations listed in the table, the specimens
are remarkably uniform in squamation and likewise in color and
markings. Neither males or females show any trace of keels on the
dorsal scales above the anal region. Variation in proportion of tail
to body length varies from twenty-five percent in a young male to
twenty-one percent in a female.

Relationship. I believe this form is a derivative of L. maculata
rather than L. mystacina, which it superficially resembles in the
broad dorsal markings. The light edging of the scales on the head,
characteristic of L. maculata, is in evidence in this species. The
species is dedicated to Dr. Hobart M. Smith, its discoverer, in recog-
nition of his extensive herpetological exploration in Mexico.

A single specimen of this species (No. 985 Balsas, Guerrero) pres-
ent in the collection of the Field Museum of Natural History, Chi-
cago, has been made available to me due to the characteristic kind-
ness of Dr. Karl P. Schmidt. This specimen is typical. Ventrals,
166; subcaudals, 68; preoculars and postoculars, 2-2; spots on body,
15; on tail, 6. The head is generally dark with the characteristic
liffht edges on the dorsal scales.

Taylor: Notes on the Genus Leptodeira 337

Leptodcira maculata (Hallowell)

(Plate XXXI, fig. 1; PI. XXXII. figs. 1-4; PI. XXXIII. figs. 1-8)

S /^//>.<a.s (iiiiiulata var. C, Dumeril and Bibron, Erp. Gen. VII, l,sr)4, \i. 1141 (two speci-
mens from Mexico).

Leptodeira annulata (part.) Giinther, Cat. Col. Snakes British Miiseuni, 1858, p. 1C6
(certain Mexican specimens).

Sibon annulata (part.) Cope, Proc. Acad. Nat. Sci. Philadelphia, ISiili. p. iiWi (Mt'xiran
specimens from Jalapa).

Megalops tnaculatus Hallowell, Proc. .\ca(l. Nat. Sci. Philadelphia, 1860, p. 488 (type
description; type locality, "Tahiti"; probably Nicaragua [fide Dunn, 1936]).

Leptodeira persanata Cope, Proc. Acad. Nat. Sci. Philadelphia, 1868 (1869), j). 310 (type
description; type locality, Mazatlan, Sinaloa, Mexico); Giinther, Biologia Centrali-.'^mericana,
Reptilia and Batrachia, May, 1895, p. 171, pi. LIV, figs. A and B (Presidio, and Mazatlan,
Sinaloa ; Santo DoiBingo de Guzman and Hda. Santa Gertrudis, Jalisco ; Mexico City ; Jalapa,
Veracruz); Bouleng-r, Cat. Snakes British Mus., Ill, 1896, pp. 93-94 (Southern Mexico
and Guatemala, several localities); Mocquard, Mission Scientifique au Mexique Rept., livr.
16. 1908, pp. 903-904.

Eteirodipsas annulata, var. scptentrionalis Jan, Icon. Gen., 39, 1872, ]^\. 1, fig. 2.

Sibon personatum Cope. Bull. U. S. Nat. Mus., No. 32, 1887, p. 67; Proc. U. S. Nat.
Mus., XIV, 1892, p. 677; Ann. Rep. U. S. Nat. Mus., 1898 (1900). p. 1107.

Leptodeira alhofusca Stejneger, Proc. U. S. Nat. Mus., 69, 1926, pp. 2-3 (refers Megalops
?naculatii>i Hallowell to this specie.s).

Leptodeira annulata annulata (part.) ,\niaral, Mem. Inst. Butantan, I\', I'Ji'j (includes
Megalop.'t marulatus as a synonym of this form).

Leptodeira septentrio7mlis maculata. Dunn, Proc. Nat. Acad. Sci., XXII. 1936, pp. 692,
697 (Cape San Lucas, Baja California; Mazatlan, Sinaloa; Zacualtipan. Hidalgo; Tuxpan,
Veracruz; "Tepanatepec" and "Zamtepec," Oaxaca:[?] Nicaragua and Costa Rica).

The type of Megalops maculatus Hallowell was supposed by
Hallowell to have been collected in Tahiti by Mr. Adams of the
Rogers Exploring Expedition. The type specimen, subsequent to its
study by Hallowell at Philadelphia, in 1860, was returned to the
National Museum at Washington and there apparently was lost or
mislaid until rediscovered by Dr. Leonhard Stejneger in 1926. After
an examination of the specimen, Doctor Stejneger identified it as be-
longing to the genus Leptodeira and the species Leptodeira albofusca
Lacepede as interpreted by Boulenger, Cat. op. cit. p. 95. Dunn [loc.
cit.\, however, regarded it as being identical with Cope's species
Leptodeira personata.

I examined this type specimen in Washington (USNM 7367).
It is in a fair state of preservation, but a serious injury to the head
makes it difficult to determine the exact characters of many of the
dorsal head scales.

The following characters can be discerned. The single preocular
is well separated from the frontal, and has a curved rather than an
angular front edge; the^ prefrontals are fused for more than half
their length, and are double the length of the internasals; loreal
elongate, much longer than high; apparently only one postocular;
fourth and fifth labials enter eye; first light band six scales long;

22—4141

338 The University Science Bulletin

no nape stripe; twenty dark bars across body; 9 + bands on the
tail; ventrals, 171; tail tip missing; the spots tend to break, and
are irregular; the contrast between the dark spots and light inter-
spaces is very pronounced; head markings indiscernible save that
the frontal appears a little darker than other dorsal head scales, and
the edges are somewhat lighter.

The scale formula, 19, 21, 21, 17, differs from the usual formula
as shown in the table.

Both Dunn and Stejneger regard it probable that this specimen
originated in Nicaragua. Dunn states that aside from six specimens
in the Museo Nacional de Costa Rica, no others are known south of
Tehuantepec.

I believe that the identity of this form with L. personata is still
open to question. Dunn advises me that data was not taken by him
on the specimens in Costa Rica, and that he does not doubt that
they originated in Costa Rica despite the lack of specific locality
data. It may be that the type of Megalops maculatn is conspecific
with L. dunckeri, since they agree in certain characters in which
both differ from Mexican forms considered here under the name of
maculata. This can only be settled by a series of specimens from
Nicaragua and Costa Rica. While I am here following Dunn in
placing Leptodeira personata as a synonym of Leptodeira maculata,
I feel that this disposition should not be regarded as final until the
form L. maculata is rediscovered in Nicaragua and Costa Rica, and
scale data taken to prove that the apparent differences, in pre- and
postoculars, and the reduced scale formulae are not sufficiently con-
stant to warrant its separation from L. personata.

Forty-five specimens in the collection have been referred to this
species. These are as follows: Nos. 5175, 5174, km. 609, Mexico-
Laredo Highway, a few miles north of Limon, June, 1936, E. H. Tay-
lor, collector; 4638-4640, Hda. La Clementina, Tamaulipas, David
Dunkle and H. M. Smith, collectors; 4631, five miles east of Jalapa,
Veracruz ; 4632, 4635, 4637, four miles east of Encero, Veracruz, July
14, 1932, and 4636, Tierra, Colorada, Veracruz, July 16, 1932,
collected by H. AI. Smith and E. H. Taylor; 5176, 15 miles west of
Veracruz, August 31, 1936, E. H. Taylor, collector; 4624-4626, near
Totolapam, Oaxaca, August 6, 1935, H. M. Smith, collector; 4643,
4644, 4651, near San Ricardo, Chiapas, September 2, 1935, and 4642,
4647-4650, near Asuncion, Chiapas, September 1, 1935, E. H. Taylor
and H. M. Smith, collectors; 4656-4657, Presidio, jMazatlan, Sinaloa,
July 19, and July 23, 1934, E. H. Taylor, collector; 4621-4623, 4641,

Taylor: Notes on the Genus Leptodeira 339

Hda. Paso del Rio, Colinia, H. M. Smith, collector; 4()45-4()4(), un-
certain Mexican locality (probably Puente Nacional, Veracruz).

Aside from the above 30 specimens are 15 specimens from Guer-
rero as follows: No. 4653 Mazatlan, 1400 meters, 5180 Agua del
Obispo 1000 m., 5182 Dos Caminos, 615 m., 4627, 4627 A, 4634, 4654
Garrapatas, 520 m., collected by E. H. Taylor and H. M. Smith,
June, ]932; Nos. 5183, 5183 A, 5184, 5185, 200 m., E. H. Taylor,
August 1, 1936; Nos. 4629, 4655, 4628, 4630, one mile north Organos,
E. H. Taylor and H. M. Smith.

These Guerrero specimens are tentatively referred to the form
Leptodeira maculata. They form two color varieties which agree
very largely in their squamation, save that one seems to have a
higher average of scale rows. In both, males show traces of keels
above anal region.

The figures (plate XXXI, fig. 1, a young specimen from Ta-
maulipas. No. 4638; plate XXXII, fig. 1, adult. No. 4628, near
Organos, Guerrero; fig. 2, adult. No. 4653, near Mazatlan, Guerrero;
fig. 3, adult. No. 4643, San Ricardo, Chiapas; fig. 4, adult, No. 4624,
Totolapam, Oaxaca) given herewith show a remarkable variation in
the color and markings. Certain of these are especially striking,
particularly the Guerrero specimen (No. 4628) which occurs to-
gether in at least two localities with the form represented by plate
XXXII, fig. 2. The differences are not due to elevation, age or sex.

The species is apparently absent on the plateau proper, but may
reach some elevation along the edges of the plateau. In the Sierra
Madre of Guerrero the highest elevation was 1,400 meters; in
Oaxaca probably less than 1,000 meters.

Certain of the differences in coloration are accompanied by cer-
tain scale differences, and differences in proportion of body to tail,
as may be shown by the following tabulation.

A. Taniaulipas specimens. Tail, 15 to 18 percent of total length; body spots,
25 to 28, often confluent; tail spots, 9 to 13; body spots reach to first
scale row (plate XXXI, fig. 1).

B. Guerrero (form with general darkened coloration tending to obscure spots) .
Tail, 19 percent of total length (?); body spots, 31 to 38; 15 spots on
tail ; body spots reach 4th to 6th scale rows. Three of eight specimens

with 25 scale rows about body, instead of 23 rows (plate XXXII, fig. 1).

C. Guerrero (form with spots distinct). Tail, 23 percent of total length {$);
29 to 37 spots on body ; tail spots, 15+ ; body spots extend to 4th scale
row (plate XXXII, fig. 2).

D. Chiapas specimens. Tail, 21 to 22 percent of total length; body spots, 28
to 30; tail spots, 10 to 14; body spots reach 1st to 4th scale rows; inter-
calated spots (plate XXXII, fig. 3).

340

The University Science Bulletin

E. Oaxaca specimens. Tail, 21 to 22 percent of total length. 25 to 29 spots
on body; 15 to 18 on tail; body spots to fourth row (plate XXXII, fig. 4).

F. Sinaloa and Colima specimens. Tail (young), 21.8 percent total length;
spots on body, 22 to 27; on tail, 8 to 10; body spots reach to the first
or second row.

With an accumulation of large series from widespread localities it

appears likely that it will be possible to define certain of these

forms clearly enough to warrant subspecific designations.

Fig. 6. Leptodeira maculata (Hallowell). EHT-HMS, No. 4624. Near
Totolapam, Oaxaca, Mexico (enlarged).

The preoculars and postoculars are invariably 2-2, save in Nos.
5180 and 4642, in which the right lower scale is fused with the
labial. The postoculars are invariably two. The upper labials are
8-8 invariably; the lower labials 10-10, save that two specimens
have 9 on one side, only, and one has 11 on botli sides. The relation
of the upper preocular to the frontal varies; in northeastern (Tam-
aulipas) specimens they are invarialjly in contact; the same is true
in all Chiapas specimens; of those from Veracruz, three specimens
have them separated; one in contact on l)otli sides, and another in
contact on one side only. Specimens from Sinaloa and Colima are
variable in this regard, some having them in contact, some separated.

Taylor: Notes on the Genl'S Leptodeira

341

some variable on the two sides of the head. In Guerrero speci-
mens, ten have them touching, three have them separated. The
temporal formula is 1 + 2 + 8. Five specimens have this varied by
fusion or by the splitting of a scale in the last series. One has the
first temporal divided vertically.

The typical juvenile coloration of the species is cream, with
blackish bars, wider on the dorsum than on the sides. The adults

Fig. 7. Leptodeira maculata (Hallowell). EHT-HMS, No. 4628. One mile
north of Organos. Guerrero, Mexico (enlarged).

have the black less intense, many, if not all, the black scales show-
ing numerous grayish flecks. The ground color becomes covered
with darker pigment of a grayish-brown color, which is usually
more intense on the outer parts of the scales. The coloration is less
intense along the borders of the dark spots and along the back. In
general, too, the head is dark, with light edges on the dorsal head
scales. One specimen from Chiapas (No. 4649) has a nuchal bar,
thus approaching the condition in rhombifera.

Three specimens from Oaxaca (4624-4626) and three from Chia-
pas 4644, 4648, 4651, have the dorsal spots reaching only to the
fourth lateral row. These likewise appear to have a greater number
of tail spots and a somewhat higher average of subcaudals.

342

The University Science Bulletin

The darker coloration of certain Guerrero specimens (plate
XXXII, fig. 1) is evident in the young specimens (Nos. 5182-5185).
(plate XXXIII, fig. 2.)

The table given herewith shows further variations in squamation
and proportions.

Table of data on Leptodeira maculata (Hallowell)

Number.

5175.
5174.
4638.
4639.
4640.

4631.
4632.
4635.
4637.
4636.

5176.
4624 .
4625.

4626 .
4643.

4647.
4642.
4650 .
4648.
4649.

4644 .
4651 .
4656.
4657.
4621.

4623
4622 .
4641
4645
4646 .

4654.
4634.

4627 .
4627A
4629.

4655 .
4653 .
4628.
4630 .
5185.

5182. .
5184. .
5180. .
5183. .
5183A

Sex
or age.

9
&

&
9

9
cf
rf'
9
9

9
9

cf
cf

d'
cf
9
&

9

yg-
yg-

9
c?
9
9
9

9
9

&
d'

yg-

9
9
9
9

9

&
9
9

Scale
formula.

21-23-17
21-23-17
21-23-17
21-23-17
21-23-17

21-23-17
21-23-16
21-23-17
21-23-17
21-23-17

23-25-17
21-23-16
21-23-17
21-23-17
21-23-17

21-23-17
21-23-17
21-23-17
21-23-16
21-23-17

21-23-16
21-23-17
19-23-17
21-23-17
21-23-17

21-23-17
21-23-17
21-23-17
21-23-17
21-23-17

21-23-17
21-23-17
19-21-15
21-23-17
21-23-17

21-23-17
21-23-15
23-25-17
23-25-17
21-25-17

21-2.3-17
21-23-16
21-23-16
21-23-17
21-23-17

Ven-
trals.

171
172
169
171
172

177
178
174
179
179

177
172
171
169
177

178
177
170
172
174

171
174
164
170
173

169
172
170
167
170

176
178
175
176
175

174
175
173
178
176

176
174
175
175
174

Sub-
caudals.

59
55
60
55
59

66

68
65

69
75
76
62

64
66
65
71

70
71
70
68

70
73
71
70
64

71
70
84

80

71 +
73

67 +

68 +
69

71
77
76
75
75

Spots,
bod.v.

26
27
26
26
27

28
26
26
25
27

25
29
29
25
29

30
30
29
28
29

30
30
24
27
21

21
24
21
24
25

37
34
33
32
34

32

29
34
38
37

31
30
33
35
34

Spots,
tail.

9
8
10
9
9

13
10
10
9
11

13t
15t
18t
11

9
11
11
14t

8t

11
12t

8
10

4t

9

14t
10
11
10

15t

13t

21

13

14t

13t
18t

17t

15t

18t

21

18t

17t

Total
length.

677
432
275
555
555

594

398*

527

490

276*

502*

405

547

529

666

492
485
495
507
400*

437
371
275
255
392*

309
530
537
630
740

487

694

592

450*

580

328
287
673
715
308

308
272
231
230
222

Tail.

120

77

53

104

104

115

101
97

88
121
115
131

83

94

105

107

97
85
60
56

66
122
118
135
146

100
137
140

137

73 +

67
132 +
137 +

66

67
65

57
52
41 +

To vent only.

Obscured posteriorly.

Taylor: Notes on the Genus Leftodeira 343

Pseudoleptodeira gen. nov.

Genotype. Hypsiglena latijaxckifa Ci\\wi\\QY=Lepto(lp'ira gvillcni
Boulengcr.

Scales in 19-21 rows; head rather broad, the snout not protruding
noticeably beyond mouth; nasal divided; loreal present; pupil ver-
tical; anal divided; scales smooth with paired apical pits; two
pairs of chinshields; tail relatively long, the subcaudals in two rows,
exceeding 60; maxillary teeth about 13-13, increasing in length
backward and followed after a short interspace by a large fang,
lacking trace of groove.

Hypsiglena discolor Giinther is tentatively referred to this genus.

Pseudoleptodeira latifasciata (Giinther)

(Plate XXXI, fig. 4)

1894. Hypsiyleiia latifasciata Giinther, Biologia Centrali-Aniericana. Rept. Batr.. Oct.,
1894, p. 138, pi. XLIX, fig. B (type description: type locality, southern Mexico): Bmilenger,
Cat. Snakes British Mus., II, 1894. p. 211: Mocquard, Mission Scientifiqiie an Mexique et
dans I'Amerique Centrale, Rept.. livr. Ki. 1908, p. 870: Dunn, Proc. Nat. Acad. Sci., 22,
No. 12, Dec. 1936. pp. 696-697 (unites H. latifasciata and Lepfodeira guilleni).

1905. Lepfodeira guiUeni Boulenger, Proc. Zcxil. Soc. London, June 6, 1905, ji. 247, pi.
\'II, fig. 2 (type description: type locality, Rio Balsas, Guerrero: Hans Ciadow, cnllector):
Mocquard, Mission Scientifique au Mexique et dans I'Amerique Centrale, livr. 16, 1908, p.
903 (description from Boulenger).

The two specimens in the collection are No. 5189, from a point
about 31/2 miles southeast of El Naranjo, Guerrero, Mexico (near
km. 190), July 18, 1936, E. H. Taylor, collector; and 4658, from
Hda. El Sabino, Michoacan, Mexico, Hobart Smith, collector.

The latter is the smaller specimen, Init it agrees in striking man-
ner with the general markings and shade of coloration of the larger
specimen. The following scale and other data are from Nos. 4658,
and 5189, respectively: Scale formula, 25, 21, 21, 17; 26, 21, 23,
17; ventrals, 188, 191; subcaudals, 80, 69; anal, 2, 2; preoculars,
2,2; postoculars, 2,2; temporals, 1 + 2 + 3 (1+2 + 4); 1 + 2 + 3;
nasal divided; upper labials, 8, 8; lower labials, 10, 10; posterior
chinshields much the longest in both; maxillary teeth, 13, 13;
ptergoid teeth, 30, 31; black bands on body, 8, 8; on tail, 3, 3.

Smith's field book gives the following data on No. 4658: "Very
large black blotches — has the parietal region of the head and the
nape of the neck brick red. It w^as found in one of the rooms of the
house at night hunting geckoes." My field book records the follow-
ing for No. 5189. "Occipital region red-orange, the light line on
lip flesh; light bands on body edged with yellow and on the median
dorsal region this yellow color is also faintly visible; the remainder

344 The University Science Bulletin

of the light blotches or bands are grayish-white. The black bands
are darkest along the median line, while the sides show some grayish
color which is washed with dim yellow; ventrals with much pigment,
the edge and the median part of the scale being the only part which
lacks pigment. The lighter color of the venter is dull cream-yellow.
The ventrals have a strong iridescence. Taken in a mass of rotten
sandstone under a vertically placed slab."

The dim nuchal line is present from parietal almost to the first
narrow bar of black. The major part of the parietals is red-orange;
the light line behind eye is strongly defined; the dark line above
connects with the first black nuchal bar; anterior labials and lower
labials with small blotches of pigment.

A recent communication from Mr. H. W. Parker, of the British
Museum, assures me that the type of Leptodeira guilleni Boulenger
does not have grooves on the fangs and is unquestionably con-
specific with Hypsiglena latifasciata Giinther.

BIBLIOGRAPHY

1929. Amau.al, Afr.\nio do. Lista remisisiva dos ophidios da regiao Neotroi)ica.
Mem. Inst. Butantan, IV, 1929, i-viii, 129-271.

1927. B.ARBOUR, Thomas, and Amaral, Afranio do. Studies on African
Ophidia. Bull. Antiv. Inst. Amer., I, No. 1, Mar., 1927, pp. 25-27.

1896. Boulenger, George A. Cat. Snakes British Museum, Vol. III. 1896,
pp. i-xiv, 1-727. pis. I-XXV.

1905. Description of new reptiles discovered in Mexico by Dr. H.

Gadow, F. R. S. Proc. Zool. See. London, 1905, Vol. II, pp. 245-247, pis. VI-VII.

1891. Cope, E. D. A critical review of the characters and variations of the
snakes of North America. Proc. U. S. Nat. Mus., XH', 1891 (1892), pp. 589-690.

1900. Crocodilians, Lizards and Snakes of North America. Ann.

Rept. U. S. Nat. Mus., 1898 (1900), pp. i-xvii, 155-1294.

1936. Dunn, E. R. Notes on North American Leptodeira. Proc. Nat.
Acad. Sci., 22, No. 12, Dec, 1936, pp. 689-698.

1858. GiiNTHER, Albert. Catalogue of Colubrine Snakes in the collection
of the British Museum. London, pp. I-XVI, 1-264.

1895. Biologia Centrali-Americana ; Reptilia and Batrachia. 1885-

1902, pp. i-xx, 1-195, Ills. 1-59. (Part dealing with Leptodeira, pp. 168-173,
published March and May, 1895.)

1908. Mocquard, F. Mission Scientifique au Mexique et dans I'Amerique
Centrale. Etudes sur les Reptiles, \\\v. 16, 1908, pp. 861-932, pis. 69-73.

1913. Werner, Franz. Neue oder seltene Reptilien und Frosche des
Naturhi.storischen Museums in Hamburg. Mitt. Nat. Mus. Hamburg, 1913,
30, pp. 1-49.

346 The University Science Bulletin

PLATE EXPLANATIONS
PLATE XXX

Fig. 1. Leptodeira punctata (Peters). EHT. No. 4614, Mazatlan, Sinaloa.

Fig. 2. Leptodeira splendida Glinther. EHT, No. 5177, 12 niile.s south,
Puente de Ixtla, Morelos.

Fig. 3. Leptodeira annulata polysticta Giinther. EHT-HMS, No. 4620,
Hda. Paso del Rio, Colima.

Fig. 4. Pseudoleptodeira latijasciata (Giinther). EHT-HMS, No. 5189,31/2
miles southeast El Naranjo, Guerrero.

Taylor: Notes on the Genus Leptodeira

347

PLATE XXX

348 The University Science Bulletin

PLATE XXXI

Fig. 1. Leptodeira maculata (Hallowell). EHT-HMS, No. 4638 (young),
Hda. La Clementina, near Forlon, Tamaiilipas.

Fi(i. 2. Leptodeira smithi sp. nov. EHT-HMS, No. 5186 (paratype), Hda.
El Sabino. near Uruapan, Michoacan.

Fig. 3. Leptodeira septentrionaUs (Kennicott). EHT-HMS, No. 4616, Hda.
La Clementina, Tamaulipas.

Fig. 4. Leptodeira bressoni sp. nov. EHT-HMS, No. 5172 (type), Hda.
El Sabino, 20 miles south, L'^iuapan, Michoacan.

Taylor: Notes on the Genus Leptodeira

349

PLATE XXXI

4
f

350 The University Science Bulletin

PLATE XXXII

Fig. 1. Lcptodcira macidntn (Hallowcll). EHT-HMS, No. 4628. one mile
north Organos, Guerrero.

Fig. 2. Leptodeira maculala (Hallowell). EHT-HMS, No. 4653, near
Mazatlan, Guerrero.

Fig. 3. Leptodeira maculala (Hallowell). EHT-HMS, No. 4643, San
Ricardo, Chiapas.

Fig. 4. Leptodeira tnacidala (Hallowell). EHT-HMS, No. 4624, Totolapam,
Oaxaca.

Taylor: Notes on the Genus Leptodeira

351

PLATE XXXII

'^^^'•^

t.^

0

%

%

3-.

^.

352 The University Science Bulletin

PLATE XXXIII

Fig. 1. Leptodeira maculata (Hallowell). EHT-HMS, No. 4653, Mazatlan,
Guerrero (young) .

Fig. 2. Leptodeira maculata (Hallowell). EHT-HMS, No. 5182, Dos
Caminos, Guerrero (young).

Fig. 3, 3a. Leptodeira maculata (Hallowell). MCZ, No. 11420, Colima,
Mexico.

Fig. 4. Leptodeira bressoni sp. nov. MCZ, 11411, (5) Colima.

Taylor: Notes on the Genus Leptodeira

353

PLATE XXXIII

¥■

23—4141

354 The University Science Bulletin

PLATE XXXIV

Fig. 1. Leptodeira bressoni sp. nov. EHT-HMS, No. 4619 (young), near
Queseria, Colima.

Fig. 2. Leptodeira annulata polysticia Giinther (young). EHT-HMS, No.
4618, Acultzingo, Veracruz.

Fig. 3. Leptodeira yucatanensis malleisi Dunn and Stuart. EHT-HMS, No.
11618 ((?), Encarnacion, Campeche.

Taylor: Notes on the Genus Leptodeira

355

PLATE XXXIV

'••

/.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXVJ June 1,1938 [No. 16

On Mexican Snakes of the Genera Trimorphodon and

Hypsigiena

EDWARD H. TAYLOR,

Departnu'iit of Ziiol(ig;\\ I'niveisity of Kansas
(Plates XXXV-XXXin, Text fig. 7)

Abstract: This study is based on specimens collected in Mexico by Edward
H. Taylor and Hobart M. Smith. The followinq; species of Trijnorphodon are
recognized: Trimorphodon bi-scatatus Dumeril and Bibron : paucnnacidatus
Taylor; lambda Cope; vilkinsonii Cope; lyrophanes (Cope); vandenburghi
Klauber; lalijascia Peters; upsilon Cope; and tau Cope.

Hyp^iqloui placed in synonymy of Lcptodcira is again validated, and recog-
nized as distmct on the basis of ungrooved back teeth, single instead of double
apical pits, together with reduced \-ertebral series, and other characters of per-
haps lesser importance.

The following forms are recognized: Hypsigiena torquata torqnnta Giinther;
torquata dunklci subsp. nov., Tamaulipas, Mexico; ajfinis Bouleuger, and
ochrorhynchus Cope.

Trimorphodon Cope

Dipsas (pt.rt.) Dumeril and Bibron, Erp. Gen., Vfl, 1854, p. 1133 (/). bi-snitata Dumeril
and Bibron).

Dipnadomorplius (part.) Giinther, Cat. Col. Snal<es British Mus.. 18.38, p. 174 (D. bi-
scutata, Dumeril and Bibron).

Trimorphodon Cope, Proc. Acad. Nat. Sci. Philadelphia, 1861, p. 297. (lyroplianes Cope).

Eteirodipsas (part.) Jan, Elenco sist. Ofid., 1863, p. 105.

Siboii (part.) Garman, Mem. Mus. Conip. Zool. Harvard College, VIH, No. 3, p. 16.

The group of back-fanged snakes belonging to this genus is char-
acterized by greatly enlarged anterior maxillary teeth followed by
smaller teeth which decrease somewhat posteriorly, and are followed
after an interspace by a pair of enlarged, grooved fangs; anterior
mandibular teeth and, to a lesser extent, the anterior palatine teeth,
enlarged; head distinct from neck; two pairs of chinshields, the an-
terior largest; seven or eight scale rows between first ventrals and
posterior lower labial; two loreals present, and frequently a third,
which is situated below the posterior; nasal divided, the nostril ver-
tically elongate; pupil elliptic; eye moderate, less than the distance

(357)

358

The University Science Bulletin

from nostril; usually three preoculars (two preoculars, one "sub-
ocular" and three postoculars) ; scales smooth (or bluntly keeled in
males), slightly oblique, with paired ai)ical pits, in 22-27 rows.
Ventrals obtusely angulate; subcaudals divided; anal divided usually
(single in vandenhiirghi) . Body compressed.

I recognized nine foiTns, which I have here treated as full species.
All save one of these are known to occur in Mexico. This exception,
vandenburghi, has been taken at the border in the southern part of
California and it is safe to state that it likewise occurs in Mexico in
northern Baja California.

Four of the species, T. vandenburghi, lyrophanes, upsilon, and bi-
scutatus, are represented by several specimens in museum collections.
The other five forms are still rare. Trimorphodon tau is probably
still known only from type; paucvmactdatus, from type only;
lambda, from three specimens; vilkinsonii, from two specimens, and
latifascia, from three. It is, of course, probable that there are mu-
seum specimens of the latter five species that have not been reported
in the literature.

Table of data on species of Tnmorphodon

Upper
labinls.

Average

ventral-subcai'.dal

total.

Scale rows.

Head markings.

bi-scvitatus (a)

9

370 (362-376)

(25-27)- (17-16)

Chevrons.

bi-sciitatus (?.)

9

338 (333-348)

(2.5-27)-( 17-16)

Chevrons.

pauciniaciilatus ....

9

329

25-17

Chevrons.

lambda

9

313 (309-317)

(22-23)-(16-15)

Chevron-'i.

vilkinsonii

9

9 (8^

308
306

23-17

(22-23)

3 black spots.

lyrophanes

Lyre-shaped mark.

vandenburghi

9 (8)

302 (299-304)

(21-23)-15

Lyre-shaped mark.

latifascia

9
8

7

293 (290-296)
288 (280-297)

25-15
(22-23)-15

Black, with red collar.

upsilon

tau

Y-shaped mark and

yellow collar.
T-shaped mark.

Trimorphodon bi-scutatus (Dumeril and Bibron)

(Plate XXV, fig. 1)

Dipsas bi-scutata Dumeril and Bibron, Erp. Gen., \'1I. 1854, p. 1153 (type description;
type locality. "Mexique").

Dipsadornorphus binrufatus Giinther, Cat. Col. Snakes British Mus., 1858, p. 176 (Mexico);
Salvin, Proc. Zocil. Soc. London, May 25, 1861, p. 228 (San Geronimo, Vera Paz, Guatemala
[27 scale rows]).

Trimorphodon hi-^rutafus Cope, Proc. .Aicad. Nat. Sci. Philadelphia, 1861, p. 297; and
Proc. Amer. Philos. Soc, XI, 1869, p. 152; Sumichrast, Arch. Sci. Phys. Natur., 46, 1873,
p. 247; and idem, pp. 253-255; Duges, La Naturak^za, VI, 1883(?), pp. 145-148, figs. 3a, 4a,

Taylor: Mexican Snakes 359

5a, (Venom); Cope, Proc. Aiiier. Philos. Soc. XXIII. 1880, p. 280; anil Proc. U. S. Nat.
Mus., XIV, 1891 (1892). p. 679; Stejneger, Ann. Rept. U. S. Nat. Mus., 1893 (1895), pp.
348, 349; Giinther, Biologia Centrali-Americana, Rept. Batr., May 1895, pp. 174-175 (part.);
Boulenger Cat. Snakes Brit. Mus., 2d Ed., 1896, pp. 54, 55 (part.); Cope, Amer. Nat., Dec,
1896, p. 1025; Cope, Ann. Rept. U. S. Nat. Mus., 1898 (1900) p. 1101; Gadow, Proc.
Zool. Soc. London, 1905, p. 224; Mocquard, Mission Scientifique au Mexique et dans
FAmerique Centrale, Rept., livr. 16, 1908, pp. 908-909; Gadow, Zool. Jahrb., Bd. 29, Heft.
6, 1910, p. 699; Werner, Mitt. Nat. Mus. Hamburg. 30, 1913, pp. 29-30; Schmidt, Field
Mus. Nat. Hist. Zool. Puhl.. XII, 1928, p. 199; Werner, Zool. Jahrb., 57, 1929, p. 181;
Aniaral, Mem. Inst. Butantan, IV, 1929, p. 201.

Eteirodipsas bisciitata Jan, Elenco Sist. Ofid., 1863, p. 105.

Trimorphodoii major Cope, Proc. Amer. Philos. Soc., XI, 1869, p. 153, and Journ. .Acad.
Nat. Sci. Philadelphia, (2), VIII, 1876, p. 131; Simiichrast, Arch. Sci. Phys. Natur., 46,
1873. p. 247; id€}n. pp. 254-256.

Sibon bisciitatum Garman, Mem. Mus. Conip. Zool. Harvard College, \'1II, No. 3, 1883,
pp. 16, 134.

Dipsas bisciitata Herrera. Cat. Coll. Rept. Batr. Mus. Nac. Mexico, 1904, p. 38 (Guana-
juato).

This species is represented in my collection by eight specimens as
follows: EHT, Nos. 5338, 5339, Hda. El Sabino, Michoacan, Ray-
mond Bresson, collector, Oct., 1935; 5145-5148, Agua del Obispo, km.
350-357, between Rincon and Cajones, Guerrero, elevation 1,000 m.,
August 1, 1936, E. H. Taylor, collector; 4588, one mile north of
Organos, Guerrero, elevation 200 m., June, 1932, E. H. Taylor, col-
lector; 4589, San Ricardo, Chiapas, September 2, 1935, H. M. Smith,
collector.

The series of specimens reported under this name by Boulenger
(1896) may very probably include more than one form. I have else-
where* suggested that the specimens from Mazatlan and Presidio,
Sinaloa, may belong to Trimorphodon paucimaculatus Taylor.

My series shows two varieties. The two Michoacan specimens
have a greatly reduced number of spots on body and tail (14-7,
14-7) as compared with those from Guerrero, which average a total
of 32 spots. They agree, however, in the very high ventral and sub-
caudal counts, the average of the total ventral-subcaudal count be-
ing 371. The single specimen from Chiapas, a male, has the ventral
count reduced, the total ventral-subcaudal count being 346. In the
specimens listed by Boulenger (1896) from Oaxaca, Guatemala (2
specimens), Panama and Central America, the range of the totals is
327 (Oaxaca) to 348 (Panama), the average, with my Chiapas speci-
men, being 339. Schmidt {loc. cit.) reports a specimen from Salva-
dor with a total count of 340. f

Males have the scaler of the posterior half or third of the body
bluntly keeled, or ridged ; they ai)pear more pronounced in the larg-
est specimens, especially above the anal region and on the tail.

* Taylor, Kansas Univ. Sci. Bull. Vol. XXIV, 1937, p. 529.

t This specimen has scale rows, 25-17, ventrals, 255, subcaudals, 85; upper labials, 7-9;
lower labials, 13-12, preoculars, 3-3; postoculars, 3-3; temporals, 3-3; length, 878 mm.; tail,
158 mm. ; ratio tail to total length, 18.

360

The University Science Bulletin

Werner (1913) re])orts a specimen 1,545 mm. long, with the scales
strongly keeled.

Table of data for Trimorphodon bi-scutatus

Number ,

Sex

Ventrals

Subcaudals

Anal

Supralabials

Infralabials

Preoculars

Postoculars

Loreals

Body spots

Tail spots

Total length

Tail

Frontal touches preoculars .

Temporals

Scales touch chirishiek's.
Scale rows t

5145

5146

5147

5148

4588

4589

5338

cf

&

cf

cf

c^

9

d^

275

274

272

272

270

261

269

101

99

99

102

100

85

101

2

2

2

2

2

2

2

9-9

9-9

9-9

9-9

9-9

9-9

9-9

11-13

13-13

13-14

12-12

14-12

10-12

13-12

3-3

3-4

3-3

3-3

3-3

3-3

3-3

3-3

3-3

3-3

3-3

3-3

3-3

3-3

2-2

3-3

2-2

2-2

2-2

3-3

3-3

19

21

20

19

24

21

14

11

10

12

13

11

10

7

653

689

575

642

440

574

938

118

128

101

116

73

90

180

no*

yes

yes

yes

yes

yes

yes

3,3,4

3,3,5

2,3,5

3,4,4

3,4,5

3.3,4

3,1,4,4

3,4,4

3,3,5

3,4,5

3,4,4

3.3,5

3,3,4

3,1,4.4

4-4

5-4

5-5

5-5

4-4

5-4

4-4

24

25

25

25

25

27

25-27

5339

&

267

95

2

9-9

1.3-13

3-3

3-3

3-3

14

7

875

166

yes

2,3,4

2,3,4

5-5

25 -27

* AliiKinnal.

t The counts are (in the above order) 38, 23, 24, 25, 16; 36, 24, 25, 23, 16; 38, 24, 25,
25, 17; 38, 25, 25, 2G, 17; 36, 25, 25, 25, 17; 36, 25, 27, 27, 17; 37, 25, 27, 25, 17; 36,

25,

17.

Trimorphodon paumnacvlatus Taylor

Trimorphodon paucimaculatus Taylor, Univ. Kansas Sci. Bull., Vol. XXIV, 1936 (Feb.
15, 1938), pp. 527-529, pi. 46, fig. 1 (type description; type locality, Mazatlan, Sinaloa.
Taylor, collector).

This species has been (iiscussed at length in the above publication.
No further specimens have been taken.

A character not previously mentioned, which differentiates this
form from bi-scutatus, is the absence of keels or ridges on the scales
in the males. This form is probably confined to the lowland coastal
region in Sinaloa, and adjoining states.

Trimorphodon lambda Cope

(Plate XXXV; fig. 4)

Trimorphodon lambda Cope, Proc. Amer. Philos. Soc, 1886, pp. 286, 287 (type descrip-
tion; type localitx-, CJiuu'nias, Sonora ; Enierich, collector); Bull. U. S. Nat. Mus., No. 32,
1887, p. 68; Proc. U. S, Nat. Mus., 14, 1891 (1892), pp. (i78-679; .Ann. Rept. U. S. Nat.
Mus., 1898 (1900), p. 1104; Taylor, Univ. Kansas Sci. Bull., 24, 1936 (1938) pp. 405-497.

Taylor: Mfa'ican Snakes 361

Trimorphodon hi-scutatus Giinther. Biologia Centrali-Auierifana. Kept, liatr.. May, 18!!;"),
pp. 174-17.') (part.); Boulenger, Cat. Snakt'.s British Mu.<;. (2), III, 189(i, p. r)4 (iiarf.).

Two specimens in my collection were discussed previously ( Taylor
op. cit.). The dental characters of a third specimen (EHT. 341A.
skeleton) follows: Four anterior maxillary teetli enlaro;ed, strongly
curved, the first more slender and shorter than the others, the third
largest of all the teeth ; the middle part of the maxillary occupied by
five teeth scarcely half the length and thickness of the preceding
teeth; this group followed by one (two on right side) large, grooved
fang, slightly curved, directed somewhat backward; seven palatine
teeth, the anterior three or four considerably larger than others;
about 15 equal ptergoid teeth; about 17 or 18 mandibular teeth, the
anterior two or three enlarged.

The relationship of this form is with Trimorphodon lyrophanes
(Cope). There are apparent differences in dentition and it is likely
that two is the normal number of loreals. Known only from south-
ern Sinaloa, Mexico. I suspect the species occurs also in Arizona,
where it has probably been confused with T. lyrophanes.

Trimorphodon vilkinsonii Cope

(Plate XXXVIII, Text. fig. 1)

Trimorphodon vilk-insonii Cope, Proc. Amer. Phil. Soc, 23. 1880. pp. 285-286 (type de-
-scription ; type locality. Chihuahua, Chihuahua; Wilkinson. coUectoi). and Proc. U. S. Nat.
I\Ius.. 14. 1891, p. 079; Giinther, Biologia Centrali-Aniericana, Rept. Batr., May, 1895, p. 174;
Cope. Amer. Naturalist, Dec, 1890, p. 1014; and Ann. Rept. U. S. Nat. Mas., 1898 (1900),
pp. 1105-1106; Crinimins, Copeia, No. 138, 1925, p. 7 (in Texas); Werner, Zool. Jahrb.. 37,
1929, p. 181; Stejneger and Barbour, Check list Amer. Amph. Rept., 3d Ed., 1933, p. 128.

Trimorphodon wilkinsonii Cope, Bull. U. S. Nat. Mus., No. 32, 1887, p. 08.

Trimorphodon upsilon Boulenger, Cat. Snakes Brit. Mus. 2d Ed. Vol. Ill, p. 55, (part.).

The type of this species is USNM No. 14268. Scale formula, 30,
21, 23, 17; upper labials, 11-11, the first five touching the anterior
chinshields, which are more than double the size of the posterior.
First neck band 5-6 scale rows wide, narrowing medially, while the
white border widens medially.

The known range of the species is Chihuahua, Mexico, and ex-
treme western Texas.

A second specimen which I have been privileged to examine is one
in the Blanchard collection, collected three miles northwest of El
Paso, close to the Rio Grande on the road to Las Cruces, June, 1935.
It presents the following characters: Rostral very much wider than
high, folding back on the snout so that part visible above is less
than one third its distance from frontal, its length about equal to
the suture between the supranasals ; greatest length of the supranasal
three fifths to three fourths the length of prefrontals; suture be-

362

The University Science Bulletin

Fig. 1. Trhnorphodoii vilkiiisonii Cope. Actual head width,

to end of parietal. 16 mm.

12 mm.; snout

tween the internasals less than half the prefrontal suture; frontal
width, 3.7 mm.; length, 5.1 mm., slightly shorter than its distance
to tip of snout (5.4 mm.) ; length of parietal, 7 mm.; parietal to tip
of snout, 10 mm.; nasal distinctly divided; three loreal scales; three
preoculars; three postoculars; upper labials, 9-9, the seventh divided
transversely on left side; lower labials, 13-13; temporals, 3 + 4 + 6;
3 + 5 + 5; five labials touch chinshields; second pair of chinshields
completely separated; about four pairs of scales between second
chinshields and first widened ventral scale; scales with double
apical pits; scale formula: 34, 22, 22, 23, 18, 16; ventrals, 228; sub-
caudals, 79; anal undivided.

Color. Brownish-gray with a series of somewhat irregularly-
shaped blackish transverse blotches; first, 9 or 10 scales back of
the parietals, somewhat narrowed on the median line, its greatest
length equal to seven scales, not reaching third scale row laterally;
this blotch followed by 27 blotches on body and tail; each blotch
is grayish in the middle, and is bordered by creamy gray; 9 blotches
on tail; forty-five pairs of ventrolateral spots touching outer scale
row, but for the most part on the ventrals, occasionally confluent
with the dorsal blotches, whicli usually terminate on second scale
row (except first blotch which is narrowest of all). The markings
on the head are diffuse, but there is evidence of a pattern with
darker areas on the prefrontals, frontal and parietals.

Taylor: Mexican Snakes 363

I owe thanks to Dr. H. K. (Uoyd for the privilege of studying
this specimen.

There is a third specimen of this rare species now in the Ameri-
can Musemn of Natural History, collected at a point "five miles
north of El Paso, Texas, on the east slope of Mt. Franklin."

Trimorphodon lyrophancs (Cope)

Lycodon lyrophanes Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 343 (type de-
scription; type locality, "Cape St. Lucas," Baja California).

Trimorphodon lyrophanes Cope, Proc. Acad. Nat. Sci. Philadelphia, 1861, p. 297 ; Proc.
Anier. Philos. Soc, 1886, p. 286; Bull. U. S. Nat. Mus., No. 1, 1875, p. 38; and idem. 32,
1887, p. 68; Proc. Amer. Philos. Soc, Apr. 1886, XXIII, No. 122, p. 286; Cope. Proc. U. S.
Nat. Mus., XIV, 1891 (1892), p. 679; Van Denburgh, Proc. Cal. Acad. Sci., (2), 5, 1895,
p. 155; Mocquard. Nouv. Arch. Mus. Hi.st. Nat., Paris, (4), I, 1899, p. 330; Cope, Ann.
Rept. U. S. Nat. Mus., 1898 (1900), p. 1102, fig. 314; Stejneger and Barbour, Check list
N. Amer. Amph. Rept., 1917, pp. 104-105,; and 2d Ed.. 1923, p. 118; and 3d Ed.. 1933, p.
127; Van Denburgh and Slevin, Proc. Cal. Acad. Sci., (4), 11, No. 4, 1921, p. 70; Schmidt,
Bull. Amer. Mus. Nat. Hist., 46, 1922, p. 697; Van Denlmrgh, Dec. Papers California Acad.
Sci., X, Vol. II, Nov. 23. 1922, pp. 884-887; Klauber. Trans. San Diego Soc. Nat. Hist.,
Vol. 5, No., 11, 1928, pp. 185-187, 190-192; Werner, Zool. .Jahrh., 37, 1929, p. 181;
Linsdale, Univ. California Publ. Zoiil., 38. No. (!. 1932. p. 383.

Klauber (1928) gives the following data on the scutellation of
this form: "Scale rows usually 22 or 23 (rarely 20, 21 or 24).
Ventrals: 223 to 243, average 232. Anal generally divided (13 di-
vided, 1 entire). Caudals, 68 to 81 pairs, average 73; supralabials,
usually 8 or 9, rarely 7 or 10; infralabials, 10-14; preoculars, 2 or 3,
normally 3; postoculars, normally 3, rarely 4. Loreals two, with a
posterior subloreal usually present. Temporals, 2+3 or 3+4, oc-
casionally 2+4 or 3+3. Body spots, 21 to 33, average 27.5; tail
spots, 10 to 14, average 12."

It will be noted that T. lyrophanes differs from lambda in having
a smaller number of spots on the tail (In lambda 17 to 18) ; a
slightly lower range of subcaudals (83-87 in lambda) and, as pointed
out under the discussion of lambda, there is considerable difference in
the teeth.

The known range is Baja California, Arizona, and very probably
also northern Sonora.

Trimorphodon vandenburghi Klauber

(Plate XXXVI, fig. 1)

Trimorphodon vandenburylii Klauber, Bull. Zoiil. Soc. San Diego, No. 1. June, 1924, pp.
27-18, fig. 3 (type descriptior^; type locality, Wildwood Ranch [elevation 1,520 feet] near
Ramona, San Diego county, California, E. B. Woodworth and L. M. Klauber, collectors) ;
Trans. San Diego Soc. Nat. Hist., V, No. 11, 1928, pp. 183-194, plates 22, 23 (Los Angeles,
Riverside, and San Diego counties. Southern California); Bull. San Diego Nat. Hist. Soc,
No. 4, 1928, p. 5; Copeia, No. 170, 1929, p. 21, (Kern and Imperial counties, California);
Bull. San Diego Nat. Hist. Soc, No. 5, 1930, p. 6; and No. 8, 1931, pp. 3, 10, 12, 16, 17,
18, 20, 23, 24, 32, 33, 35, 44, 45, 51, 52, 62, 72: and No. 11, 1934, pp. 20, 21; Stejneger
and Barbour, Check list N. Amer. Amph. Rept., 3d Ed., 1933, p. 128 (Kern, Imperial, Inyo
and San Diego counties, California).

364 The University Science Bulletin

I collected a single specimen (Kansas University Mus. 8497) of
this species from under a flake, pried from a granite boulder a few
miles east of San Diego, Cal., September 1, 1928. The specimen
agrees with the type description in all pertinent characters. The
proximity of this and other records to the Mexican border (Dulzura,
San Diego county, from which Klauber [1928] records a specimen,
is also near the border) suggests that the species is a habitant of the
northern part of Baja California.

Ventrals, 239; anal entire; subcaudals, 65; upper labials, 9-9;
lower labials, 12-13, five touching anterior chinshields; preoculars,
postoculars and loreals, 3-3; head (mm.), 13.5X11; frontal shorter
than its distance to end of snout; 40 blotches on body, 16 on tail;
total length, 485; tail, 72.

The undivided anal scute (rarely divided) will separate this form
from its congeners in most cases.

Trimorpliodon latijascia (Peters)

(Plate XXXVI, fig. 2)

Trimorpliodon biscutatus var. latifascia Peters, Monatsb. Akad. Wiss. Berlin, 1869, p.
877 (tjpe description; type locality, Puelila, Mexico; Doctor Berkenbusch, collector).

Trimorpliodon coUaris Cope, Journ. Acad. Sci. Philadelphia, (2), VIII, 1876, p. 13 (type
description; type locality, Orizaba, Veracruz: Doctor Suniichrast, collector); and Proc. U. S.
Nat. Mus., XIV, 1891, p. 679.

? Sibon biscutatum var. latifasciatum Garman, Mem. Mus. Comp. Zool. VIII, No. 3, 1883,
p. 16-17.

Trimorphodon itpsiloii Boulenger, Cat. Snakes British Mus. (2), III, pp. 55, 56 (part.);
Cope, .\nn. Kept. U. S. Nat. Mus. 1898 (1900), pp. 1104-1105 (part.).

The discovery of a specimen of Trimorphodon, related to, but dis-
tinct from Trimorphodon upsilon necessitates the revival of Peters'
name latifascia. Cope described a species, Trimorphodon collaris
seven years after latifascia was described, but it seems to be a syn-
onym. My specimen was obtained twelve miles south of Puente
de Ixtla, Morelos.

Peters' description, though brief, offers the following data: Scales
in 21 to 22 rows; ventrals, 206 to 210; 13 to 14 broad, transverse
bands on body; 5 to 6 on the tail. Bands gray-brown, edged with
black. No distinct V-shaped markings on the head ; a bright neck-
band, which, upon the parietals, forms a V-shaped indentation and
covers the temporal region and three posterior supralabials.

My specimen (EHT Xo. 5439) and the type of Trimorphodon
collaris Cope (USNM No. 26499 ( have the following characteristics,
respectively: Ventrals, 218, 211; subcaudals, 78, 79; upper labials,
9-9, 9-9; lower labials, 12-13, 12-12; preoculars, 3-3, 3-3; post-
oculars, 3-3, 3-3; loreals, 3-2, 3-3; scale formula, 33, 25, 23, 23, 15:

Taylor: Mexican Snakes 865

31, 25, 23, 23, 15; scales touching chinsiiields, 5-(i, 4-5; anal divided
in both; temporals, 3, 4, 5; (3, 3, 4, 5), and 3, 4, 5; body spots, 15,
16; tail spots, 8, 8. Total length, 256 mm.; tail, 42 mm.

]\Iy specimen presents the following additional characters: 8th
labial does not reach the lij); rostral very short, about one fifth its
distance from the frontal; frontal one fourth longer than its distance
from end of snout, and one sixth shorter than the parietals (about
as long in type of collaris) ; total length, 256 mm.; tail length, 42
mm.; frontal, 3.5 mm.X2.7 mm.; head, 11.1 mm.X7 mm.; eye, 2.5
mm.; eye to nostril, 2.5 mm. The frontal not in contact with pre-
ocular (in contact in ty])e of collaris).

The bands on the back are wide, the four anterior on back cov-
ering 19, 15, 16, 16' scales, respectively, the intervening red spaces
3I/2 or 3 scales wide on median line; the black bands are much nar-
rowed below, the first four covering 8, 9, 9, 7 ventrals while the in-
tervening reddish color covers 10, 9, 9, 10 ventrals; posteriorly the
dark bands become narrowed, covering four or five ventrals; the
reddish areas cover 12 ventrals. However, throughout the body
there are intercalated dark spots tending to divide the red areas,
crossing ventrals and extending up to fourth lateral scale row; the
black bands have a trace of a white line dividing the spots trans-
versely.

The top of the head is black, less dense on the tip of snout; the
first red collar is about four scales wide medially, tending to en-
croach on the parietals; last three labials red, the fourth to sixth
with red spots (reddish areas probably brown in adults).

Tnmorphodon upsilon Cope

(Plate XXXV, fig. 2)

Trimorphodon upsilon Cope, Proc. Anu-r. Philos. Soc, XI, 1869, p. 1.52 (type description;
type locality, "Guadalaxara" West Mexico, I. I. Major, C), and idem, XXIII, 1886, p.
286; Bull. U. S. Nat. Mus., No. 32, 1887, p. 68 (Batopilas, Chihuahua; Guanajuato;
Zacualtipan, Hidalgo); Cope, Proc. U. S. Nat. Mus., XIV, 1891, p. 678; CJiinther. Biologia
Centrali-Aniericana, Kept., May 1895, p. 175 (Ventanas, Durango; Guanajuato; La Cuuihre
de los Arrastrados, Jalisco [?]; Jalapa, ^'erac^uz); Boulenger, Cat. Snakes Brit. Mus., 2d
Ed, III, 1896, p. 54; Cope, Ann. Rapt. U. S. Nat. Mus., 1898 (1900), pp. 1104-1105, fig.
315; Mocquard, Bull. Soc. Phil. Paris (9). I. No. 4, 1899, p. 157 ("Sierra del Nayarit");
Gadow, Proc. Zool. Soc. London, June 6, 1905, pp. 224, 231, 233 (distribution); Mocquard,
Mission Scientifique au Mexiciue et dans I'Amerique Ccntrale, Rept., livr. 16, 1908, pp. 910-
911, pi. 74. figs. 2, 2a, 2b; Gadow, Zoiil. Jahrb.. Bd. 29, Heft. 6, 1910, pp. 666. (i!»7. 761,
702 (ranges in elevation from about 1,000 feet to 7.(100 feet); Amaral. Mem. Inst. Butantan,
IV, 1929, p. 202.

Eteirodipsas biscutata Jan, Icon Ofid. XXXIX, 1872, pi. I, fig. 3.

Sihnii upsilon Garman. Mem. Mus. Comp. Z>>ol., VII. No. 3. 1883. Ji. 134.

A single specimen (EHT, No. 4569; length, 656; tail, 130 mm.)
of this species was taken by Dr. Hobart Smith near ]\iagdalena,
Jalisco, June 17, 1935, at an elevation of 1,300 m. 1 have also

366

The University Science Bulletin

examined several specimens of this form in the United States Na-
tional Museum. Several specimens lack locality data. These are
USNM. Nos. 9912 (2 specimens), 25361, 26138, and 26139. They
were collected by Duges and are probably from Guanajuato. Nos.
21419 and 31358, Type, Guadalajara, Jal; No. 46334, San Juan
Capistrano, Zacatecas.

The range of the form is chiefiy in western Mexico from Chi-
huahua south through Durango, Guanajuato, Jalisco and Nayarit.
Cope's (1887) specimen from Zacualtipan, Hidalgo, and Giinther's
(1895) from Jalapa, Veracruz, are the only records for the eastern
side of the plateau.

The following tabic includes data taken from certain of these
specimens.

Table of data on Trimorphodon upsilon Cope

Number

9912

9912.\

12419

25361

26138

4569

31358

46334

Museum

USNM

USNM

USNM

USNM

USNM

EHT

USNM

USNM

Sex

&

9

7

9 (?)

9

cf

cf

9

\'eiitral?^ ... .

217

69

3-3

3-3

228

59

3-3

3-3

218

62

3-3

3-3

223

66

3-3

3-3

211

68

.3-3

3-3

222

75

3-3

3-3

222

73

3-3

3-3

232

61

Preoculars

3-3

Postoculais

3-3

Loreals

3-3

3-3

2-2

3-2

2-2

2-2

3-3

3-3

Upper labials

8-8

8-8

8-8

8-8

8-8

8-8

8-8

9-9

Lower labials

12-12

13-13

12-12

11-12

12-12

12-11

11-11

11-13

Scales touch
chinshields

4-4

4-4

4-4
3,4,5

4-5
3,3,3

.5-5

3,3,4

2,4,5

Bands, body

30

28

23

31

28

25

24

29

Bands, tail

15

11

11

13

15

13

13

11

Scale formulae, No. 9912, 31, 22, 22. 21, 1."); 9912A, 30. 23, 23, 24, 10; 26138. 32. 24,
22, 22, 10; 4569, 30, 22, 22, 17. 15.

Trimorphodon tau Cope

Trimorphodon tau Cojie, Proc. Anier. Philos. Soc, 9, 1869. p. 152 (type descriptitn;
type locality, Isthmus of Tehuantepec, Mexico); and idem. Vol. 23, 1886, p. 286; Bull.
U. S. Nat. Mus., No. 32, 1887, p. 68; Proc. U. S. Nat. Mus., 14, 1891, p. 078 (key);
Giinther, Biologia Centrali-Americana, Rept. Batr., May 1895, p. 174; Boulenger, Cat.
Snakes Brit. Mus., 2d Ed. 1896, p. 50; Cope, Ann. Rept. U. S. Nat. Mus. 1898 (1900), p.
1101; (?) Mocquard, Bull. Soc. Phil. Paris (9), I, No. 4, 1899, p. 157 (.Jalisco, Diguet,
collector); Mocquard, Mission Scientifique au Mexique et dans I'.Auierique Centrale, Rept., livr.
16, 1908, p. 912; Gadow, Proc. Zool. Soc. London, 190;-), p. 224; .\niaral, Mem. Inst.
Butantan, IV, 1929, p. 202.

The type of the species is probably the only known S]iccimcn.
Mocquard (1899) lists a specimen from Jalisco, but later (Moc-

Taylor: Mexican Sxakes 367

qiiard, 1908) states that the type is the only known specimen.
Whether the specimen Usted in 1899 is lost or has received another
designation I cannot say.

I have examined the type (USNM. No. 30338, Tehuantcpec,
Sumichrast, collector). It is in rather bad condition, being much
softened. The striking head pattern is still quite evident. The scale
formula is 31, 21, 23, 15. It is known definitely only from the type
locality. The following data are given in the type description:

Scales in twenty-three series. Muzzle projecting considerably be-
yond the mouth. Rostral plate somewhat produced behind; in-
ternasals about one fourth size of prefrontals, which are as long as
wide. Frontal with straight lateral margins. Parietals not longer
than frontal, regularly rounded behind. Nostril in middle of nasal.
Three loreals, three postoculars, three preoculars. Temporals, 2+ 3
+ 4; six upper labials, the fifth probably composed of two plates
fused, as it is twice as long as deep on both sides. The fourth and
fifth enter the orbit, the third is cut down by the lower loreal and
preocular. Lower labials, eleven. Body strongly compressed. Total
length, 236; tail, 35.

Above gray with twenty-three jet-black rhombs, which extend to
the ventrals by their lateral angles. Tail with ten rhumbs; sides of
belly black spotted. Head gray with a black mark above as far as
the middle of the parietals, but with two, lateral, ear-shaped projec-
tions on the same; a pale T-shaped mark extends transversely be-
tween the orbits, and longitudinally to the end of the muzzle.

Hypsiglena Cope

Leptndeira (part.) Giinther, Ann. Mag. Nat. Hist. (3), V. 1860, p. 170.
Hypsiglena Cope, Proc. Acad. Nat. Sci. Philadelphia, .June, 1860, p. 24<j (generic de-
scription; type ochrorynchus).

Pseudodipms Peters, Monatsb. Akad. Wiss. BerUn, 1860, p. 521.
Comastes Jan, Elenco Sist. Ofid., 1863, p. 102.

Body small, slender, somewhat cylindrical; head distinct from
body, the snout projecting beyond mouth; head plates normal; two
nasals distinct or united above nostril; two lor three I preoculars,
two postoculars; loreal present. Temporals, normally 1-2; scales
smooth (save on sides above anus in males), in 19-21 nnvs; apical
pits single; anal plate divided; eye very small. i)upil vertically
elliptic; tail very short, less than one fourth body length; ventrals
not angulate; subcaudals divided; anterior maxillary teeth four to
eight, subequal, followed after a space by one or two large, un-
grooved, fanglike teeth.

I am fully convinced that the species here considered under this

368 The University Science Bulletin

genus, H. torquata torquata, H. t. dunklei subsp. nov., H. affijiis,
and H. ochrorhynchus, form a generic group related to Leptodcira,
but differing in having the snout projecting more, a proportionally
smaller eye, a much shortened tail, usually less than 10 maxillary
teeth, single apical pits, posterior fangs lacking grooves; a smaller
number of subcaudal scales. I have not regarded Leptodeira guil-
leni Boulenger (Hypsiglena latifasciata Giinther) or Leptodeira dis-
color Giinther as belonging in the genus Hypsiglena.

Giinther ((k't., 1894), loc. cit., confused his specimens of Hypsi-
glena, throwing together as one species all the species here recognized
(i. e., torquata, ochrochynchiis, and affinis). Boulenger (1894) on
the other hand, using the same specimens available to Giinther,
recognized the three species.

With accumulation of more material from western JNIexico it will
be possible, no doubt, to demonstrate that certain forms included
under torquata torquata are worthy of subspecific designation.

Key to the Forms of Hypsiglen.\

A. Scales in 19 rows around niifidle of l)o(ly: a single preocular; upper labials 7, lower
labials 10; ventral-subcauilai count 207-213; head narrow, with about 23 scales

about posterior part of head; a light nuchal band H. affinis Bouleng r,

AA. Scales in 21 i-ows around middle of body; two preoculars (1 preocular, one sub-
ocular) ; upper labials 7 or 8 ;
B. No light neck band ; first nuchal spot single (rarely divided into two or three

parts) connecting with dark lines behind eyes H. ochrorhytichus Cope,

BB. A light nuchal band, not or rarely interrupted medially or lat rally, sometimes

involving part of parietal region.

C. Rostral large, pushing far ui between jirefrontals, tlie jiart visilile aljout

equal to three fourths the distance lietween rostral and frontal; head

broad, 30 scales about posterior part of head; 8 upper labials; 11 lower

labials; knobs or keels on lateral scales abo\e anus in males; 162

ventrals ; .57 sulicaudals (male) //. torquata tlioiklei subsp. nov,

CC. Rostral not pushing between the prefrontals or but slightly: part visible
above eciual to one half or less of the distance between the rostral and
frontal; 2.5 or less scales about back part of head; upper labials, 7 or
8; lower labials, 9 or 10; aiiiiartntly no scales with keels or knobs on
sides above anus in males; ventrals. 104-174; subcaudals, 36-56

H. torquata torquata Giinther.

Hypsiglena ochrorhynchus Cope

Hi/psit/leiia onliror/iyiiclius Cope, Proc. Acad. Nat. Sci. Philadelphia, 12, Nov. 15, 1860,
p. 246 (type de,scription ; type locality, "Cape St. Lucas," Baja California, John Xantus,
collector); and Bull. U. S. Nat. Mus., No. I, 1875, p. 38; Lockington, Amer. Nat., XIV, 1880,
p. 295; Yarrow, Bull. U. S. Nat. Mus., No. 24, 1883, pp. 15, 97 (Cape San Lucas, La Paz,
Baja California; Durango, Mexico); Garman, Mem. Comp. Zool., Harvard College, XUi.
No. 3, 1883, pp. 80, 161; Cragin, Bull. Washburn College Lab. Nat. Hi.st., I, 1884, p. 8
(Guaymas, Mex.); Cope, Proc. Amer. Pliilos. Soc. Philadelphia, XXIII, p. 285 (Chihuahua);
and Bull. U. S. Nat. Mus., No. 32, 1887, p. 78; fielding. West Amer. Scirnt., Ill, No. 24,
1887, p. 98; Cope, Proc. U. S. Nat. Mus., XIV, 1891 (1892), p. 617; Steneger, N. Amer.
Fauna, No. 7, May, 1893, pp. 204, 205 (Cape St. Lucas); Boulenger, Cat. Snakes British
Mus., Vol. II, 1894, p. 209; Van Denburgh, Proc. Cal. .Acad. Sci. (2), Vol. 5, 1895, p. 145;
Occ. Papers Cal. Acad. Sci., V, 1897, p. 178; Mocquard, Nmiv. Arch. Mus. Hist. Nat.,
Paris, (4), I, 1S!»;». p. 325 (Mulege, Baja California); Cope, Ann. Ript. U. S. Nat. Mus.,

Taylor: Mexican Snakes 369

1898 (inOO), 10. 953, fig. 245; Brown, Pior. Acad. Nat. Sci., Philadelphia, 1001, p. 87; Van
Denburgh, Proc. Cal. Acad. Sci. (3), ZoGl., \'(>1. 4, No. 5, 1906, p. 65; Ditmars' Reptile
Book, 1907. p. 329, pi. CI, fig. 1; Giinnell, Univ. Cal. Publ. Zoo!., Vol. 5, No. 1, 1908, p.
165: Van Dcnhuich and Sleviu, Proc. California Acad. Sci. (4), Vol. 3, 1913, p. 414; Atsatt,-
Univ. California Publ. Zool., 12, No. 3. 1013, p. 42; Riithling, Copeia, No. 15, 1915; Van
Denburgh and Slevin, Proc. California Acail. Sci., (4), 5, 1915, p. 106; Grinnell and Camp,
Univ. California Publ. Zool., 17, No. 10, p. 188; Bentley, Copeia, No. 61, 1918, p. 83;
Cowles. Jourii. Ent. Zoill. Pomona College, XII, 3, 1920, 66; Stevens, Trans. .San Diego Soc.
Nat. Hist. Ill, 4, 1921, p. 61; Nelson, Mem. Nat. Acad. Sci., XV'I, 1921, pp. 114, 1151
Klauber, Bull. Zool. Soc. San Diego, No. 8, 1931, p. 71, and No. 9, 1932, pp. 25, 80; and
Copeia, Oct. 7, 1932, No. 3, p. 126; Linsdale, Univ. California Publ. Zool., 38, No. 6, 1932.
p. 380 (San Ignacio, Coniondu and Eureka in Baja California); Stejneger and Barbour,
Check list N. Amer. Amph. Rept., 3d Ed., 1033, p. 113; Allen, Occ. Papers Mus. Zool.
Univ. Mich., No. 259, p. 12 (Hermosillo, Son.); Klauber, Bull. Zool. Soc. San Diego, No. 11,
1934, p. 19; Taylor, Univ. Kansas Sci. Bull.. 24, 1936 (Feb. 15, 1938), pp. 494-495.

Hypsiylena chlorophaea. Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 247 (type
description; type locality. Fort Buchanan, Arizona, Irwin, collector); Stejneger, N. Amer.
Fauna, No. 7, 1893, p. 205; Mocquard, Nouv. Arch. Mus. Hist. Nat., Paris, (4), I, 1899, p.
325, and Mission Scientifique au Mexique et dans I'Amerique Centrale, livr. 16, 1908, p. 869,
pi. 69, fig. 1. la-d; Gannan, Bull. Essex Inst.. 16, Jan. 9, 18S4, p. 30.

Hypsifileiia ochrorhynchus chlorophaea Cope, Proc. Acad. Nat. Sci. Philadelphia, 1866, p.
304; and Bull. U. S. Nat. Mus., No. I. 1875, p. 38 (Arizona); Coues. Wheeler's Rept.
Surv. W. lOOth Merid.. V, 1875, p. 622; Yarrow, Bull. U. S. Nat. Mus. No. 24, 1883, pp.
15, 97, 190; Garman, Mem. Mus. Comp. Zool., Harvard College, VIII. No. 3, 1883, pp. 80,
161; and Bull. Essex Inst., 16. 1884, p. 30.

Hypsirjleita torqiiata Biologia Centrali-Aniericana. Rept. Batr., Oct., 1894, p. 137 (part.);
Mocquard, Mission Scientificjue au Mexicpie et dans I'Anierique Centrale, Rept., livr. 16. 1908,
pp. 866-868, pi. 69, figs. 3, 3a. 3c, 3d (part.) (He .states, Bocourt believed //. nrhmrhi/iichus
distinct from H. torquata).

Hypsiglena ochrorliytichiis ochrorhyurhii.i Stejneger and Barbour, Check list N. Amer.
Amph. Rept., 1917, p. 93; Van Denburgh and Slevin, Proc. California Acad. Sci., (4), XI,
1921, pp. 28, 52, 68; Schmidt, Bull. Amer. Mus. Nat. Hist., XLVI, Dec. 7, 1922, p. 692;
Klauber, Bvdl. Zool. Soc. San Diego, No. 1. .June 1, 1925, p. 16; Van Denburgh, Occ. Papers
California Acatl. Sci., X, 2, pp. 780. 783, pi. Sii; Stejneger and Barbour, Check list N.
Amer. Amph. Rept., 2d Ed., 1923, p. 104.

Leptodeira torquata venusta Dunn, Proc. Nat. Acad. Sci. 22. 1936, pp. 691, 695.

Leptodeira torquata ochrorhynchus Dunn, Proc. Nat. Acad. Sci. 22, 1936, pp. 691, 695.

Hypsiglena texana Stejneger N. Amer. Fauna., 7, May, 1803, p. 205 (type description;
type locality, "between Laredo and Camargo. Texas); Strecker, Baylor Bull.. XVII, No. 4,
1915, p. 40.

Hypsiglena ochrorhynchus texana Stejneger and Barliour, Check list N. Amer. .\uiph.
Rept., 1017, p. 93; 2d Ed. 1923. p. 104.

I have hesitated to venture an opinion on the validity of the sup-
posed species, Hypsiglena venvsta Mocquard, H ypsiglena chloro-
phaea Cope or Hypsiglena texana Stejneger, because of insufficient
specimens. It is likely that when sufficient material is available,
certain of these will be recognized as subspecific forms. (Dunn,
1936, has recently recognized venusta.) Differences in the character
of the skull is marked in specimens from San Diego county. Cali-
fornia, Northern Mexico and Arizona. The snout is short and blunt
in the former, elongate and projecting over the mouth in the latter.

The species is represented in my collection by six Mexican speci-
mens, from the following localities: EHT, No. 4595, Vo miles north-
west of SaUillo, Coah., August 24, 1932, H. M. Smith; 4596. five
miles southwest of Hermosillo, Sonora, June 22, 1934; 4597, near La
24—4141

370

The University Science Bulletin

Posa, 10 miles northwest of Guaymas, Sonora, June 20, 1934; 4598,
five miles northwest of Guaymas, Sonera, 1934; 4599, 32 miles west
of San Pedro, Coah., August 25, H. M. Smith; 5202, Huasteca
Canon, 11 miles w^est of Monterey, Nuevo Leon, June 16, 1936.
(Unless otherwise mentioned, the specimens were collected by me.)
The scale formula for the three Sonora specimens is 21, 21, 17, 15;
that for the three specimens from Nuevo Leon and Coahuila is 21,
21, 19, 17.

Table of data and measurements (in mm.) of Hypsiglena ochrorhynchus Cope.

Number .

Sex

Ventrals

Subcaudals. . . .
Supralabials . .
InfralabiaLs . . . .

Preoculars

Postoculars . . . .
Temporals . . . .

Anal

Total length...

Tail

Frontal length.
Frontal width .
Head length. . .
Head width . . .

Eye

Eye to nostril .
Spots

4597*

177

54

8-8

10-10

2-2

2-2

1-2-3

2

408

G9

4

2.5
17
10
2.3
2.8
59?

4598

181
63
8-7
10-10
2-2
2-2
1-2-3
2
397
73
3.9
2.3
16
9.6
2.3
2.95
70

4596

178
60
8-8
10-10
2-2
2-2
1-2-3
2
399
74
4.2
2.7
15.8
9

2.3
2.8
70

4595

9

175

41

8-8

11-11

2-2

2-2

1-2-3

2

365

53
3.6
2.8

15
9.2
2.15
3

53

4599

174

56

8-8

10-10

2-2

2-1

1-2-3

2

362

66

4

2.3
13.5
9

2.2
2.8
49

5202

9

172

41

8-8

10-10

2-2

2-2

1-2-3

2

.391

55
3.8
2.8

13.7
9.3
2.2
2.6

56

* The numbers 4590. 4507, 4508 bore the field numbers of 120, 266, 281, respectively, and
are so referred to in Taylor (1036). The total length as given is in ermr. These are
snout-to-vent measurements. No. 281 should read, 326 mm.

Hyimglena affinis Boulenger

(Plate XXXVII, fig. 3)

Hypsiglena torquata GUnther, Biologia Centrali-.\mericana, Kept. Batr., Oct. 1894, p. 137
{part.).

Hypsialena affinis Boulenger, Cat. Snakes British Mus., II, 1894, pp. 210, 211, pi. 8 (type
description : type locality Zacatecas and Jalisco) ; Mocquard, Bull. Soc. Phil. Paris, (9), I,
No. 4, 1899, p. 157 (Guadalajara); and Mission Scientifique au Mexique et dans I'Amerique
Centrale, Kept. livr. 16, 10(i8, pp. 868-8(10, pi. 60, fig.s. 2, 2a, 2b, 2c (Mexico); Werner, Zool.
Jahrb, 57, 1920, pp. 124, 125 (Key).

Leptodeira torquatus torquatus Dunn., Proc. Nat. Acad. Sci., 22, 1936 (part.).

This species is represented in the collection by a single male
specimen, No. 4601, collected by H. M. Smith, near Magdalena,
Jalisco, June 1, 1935. It agrees generally with the type description.

Taylor: Mexican Snakes 371

Scale formula, 23, 19, 19, 17, 15; vontrals, l(i2; subcaudals, 45;
anal divided; upper labials., 7-7; lower labials, 10-10; ])reoeulars,
1-1 (no subocular) ; postoculars, 2-2; temporals, 1 + 2 + 3; 5 labials
touch anterior chinshields, whicli are very distinctly larger than the
posterior; scale pits single; order of size of labials, 1, 2, 3, 4, 7, 6, 5.
Total length (mm.). 297; tail, 4(5; frontal, 3 X 2.7; head length, 10;
width, 7.6; eye, 1.7; eye to nostril, 2; maxillary teeth, 4, with four
interspaces (whicli probably bear teeth normally) followed, after a
short diastema, by a pair of enlarged fanglike, ungrooved teeth.

The general ground color is gray-brown with a median series of
spots and three lateral series on each side, the median spots largest,
but some tending to break in two on the median line. The smaller
lateral spots are arranged to alternate with the median spots and
with each other. The head is brownish, and a narrow longitudinal
nuchal line is present. The nuchal collar is cream color. The first
dark nuchal blotch covers six to eight scale rows; the cream collar,
which jirecedes it, covers six scale rows. The dark band running
back from the eye does not cross the cream collar to join the dark
nuchal blotch laterally. The band is bordered by a narrow white
line above and below. Very slight pigmentation is evident on the
edges of the chin. Below, the abdomen and subcaudal regions are
white. On the sides above the anus, the scales bear rounded,
knoblike tubercles or keels (probably present only in males).

I am convinced that this form is distinct from either H. torquata
or H. ochrorynchus, on the basis of the presence of only 19 scale
rows, the absence of the lower preocular (subocular) and the
characteristic color pattern.

The present known distribution includes Zacatecas, and Jalisco
in IMexico.

Hypsiglena torquata torquata Giinther

(Plate XXXMI. tiff. 3)

Leptodeira torquata CUiather, Ann. Mag. Nat. Hist. (3), V, Feb. 1860, p. 17(i, pi. 10,
fig. A (type description: type locality, Lagnna I., Nicaragua); Trooschel, in Miiller, Reisen
in iltn \'erein;gten Staaten. Canada und Mexico, UI. ISfif). p. 612.

Pscudodipsas jallax Peters, Monatsb. Akad. Wiss. Bi-rlin, 1860, p. .520.

Liophis jarril Duges, Mem. Ac. Montpelier, \'I, 18()6, proc.-Verb. p. 32 (fidi Roulenger).

Cotnastes quincnnciatus Jan, Elenco Sist. C)fid.. 1863, )). 102, and Icon. Gm. Ofid., 38,
1S71. pi. 1, fig. 1. ("Mazatlan," "Co-sta Rica," "Caracas," "Mexico"); Trooschel, in Miiller,
Reisen in den Vereinigten Staaten, Canada und Mexico, III, 1865, p. 612.

Hypsiglena torquata Cope. Bull. U. S. N. M., No. 32, 1887, i). 78: Ciintlier, Biologia
Centrali-.\niericana, Rept. Batr., Oct.. 189.5, ]i. 137 (part.); Boulenger, Cat. Snakts British
Mus., n, 1894, p. 210, 359 (Ventanas, Durango ; Presidio, near Mazatlan, Sin.; Nicaragua);
Gadow, Proc. Zool. Soc. London, June 6, 1905, pp. 224, 241 ; Mocquard, Mission Scientifique
au Mexique et dans I'-Amerique Centrale, Reptiles, li\r. 16. UH.iS, |i|i. 867, 868 (part.) figs.
3 (?); Wern»r, Zool. Jahrb., 57, 1929, pp. 124, 125.

372 The University Science Bulletin

Lrptodeira torquata torquata Dunn, Pioc. Nat. Acad. Sci. 22, 193G, pp. G91, 694, 695
(part.)

This species is represented by two specimens collected twelve
miles south of Puente de Ixtla, Morelos. Both were collected
from under rocks. The following data arc taken from EHT.
Nos. 5200, 5201, respectively (measurements in mm.) ; sex, 5 ,
$ ; scale formula 25, 21, 21, 19, 17 in both; ventrals, 166, 164;
subcaudals, 39, 36; upper labials, 7-8, 7-7; lower labials, 9-10, 9-9,
the first pair separated; preoculars (pre- and subocular), 2-2, 2-2;
postoculars, 2-2, 2-2; temporals, 1 + 2 + 3. 1 + 2 + 3 (2 + 2 + 3
on right side); scales touching first chinshields, 4-5, 4-4; total
length, 460, 412; tail length, 59, 54; scale pits single; anterior
chinshields largest or equal in size to second pair; white collar, 4
scale rows wide; frontal longer than distance to end of snout in
both; frontal length, 4.2, 4; frontal width, 3.1, 2.9; head length,
15.3, 14.3; head width, 11, 10; eye length, 2.2, 2; eye to nostril, 3,
2.8; spots on dorsal line approximately, 56, 52. The anterior
maxillary teeth are 4 on each side (with probably four missing
teeth, making a total of eight on each side), followed by two large
fangs, which show no trace of a groove. There is a nuchal dark line
in one, only small spots in the other, neither connecting with the
large dark nuchal blotch. This blotch is four scale rows long medi-
ally, seven or six and one lialf long laterally. The dorsal pattern of
both specimens consists of a median series of spots which are divided
in two at one or two regions on the back; when unbroken the spots
are quadrangular and separated by transverse cream lines. The
spots are brownish lavender on a dull, gray-lavender; two rows of
smaller dots laterally; top of head dark with minute whitish flecks;
a diagonal cream-white line from below tlie eye and a white line
along the lower labials; a broad brown band from eye to jaw angle,
bordered above with cream; chin heavily pigmented; outer edges of
the ventrals with some pigment.

USNM No. 31385 j , Colima, Mexico. Xantus coll. This speci-
men shows certain anomalies. The prefrontals fused; posterior
chinshield longer tlian anterior; 4 scales touch anterior chinshields;
upper labials, 8-7; lower labial, 8-8; 2 preoculars; 2 postoculars;
rostral ])ushing between the internasals slightly. One very large
anterior temporal on left side, with a tiny scale below (destroyed
on left side); frontal, length, width 3.5 mm. X 2.7 nun.; parietal,
4.5 mm. long; parietal to end of snout, 5 mm.; nasal divided; eye
diameter, 1.7 mm.; eye to nostril, 2 nun.; scales preceding divided

Taylor: Mexican Snakes 373

preanal, partially tu^ed; scale luriuula, 22-21-21-17-15; ventrals,
166; subcaudals, 42. Typical markings; slight amount of pigment
on anterior chinshields; slight pigment on edges of ventrals; about
52 median dorsal spots. Head flecked with deep brown.

USNM No. 51479 5 , San Bias (?Nayarit), Mexico. Prefrontals
normal; rostral short above, not pushing between internasal; poste-
rior chinshields longer than first; two preoculars; two postoculars;
temporals, 1 + 2 + 3. frontal length, width 3X2 mm.; frontal to
tip of snout, 2.7 mm.; parietal, 4.2 mm.; upper labials, 8-8; lower
labials, 10-10; scale formula, 26-21-21-19-17; ventrals, 171; sub-
caudals, 39. Anal divided. Five scales touch first chinshields, which
are shorter than posterior pair. First neck band notched behind,
7 or 8 scale rows wide; 3-4 scale rows behind parietal, white; head
flecked with deep brown; 54 dorsal spots. Total length, 295 mm.;
tail, 40 mm.; head length, 12 mm.; width, 6.5 mm.

USNM 46513. Tupataro, Midioacan, has a band on neck with
a narrow median and two lateral projections pushing forward, the
lateral ones connecting with the line behind eye; head light, finely
flecked to between eyes. This marking on neck is strongly sug-
gestive of the nuchal mark in H. ochrorhyuchus.

The parietals are short (5.3 mm.) ; parietal to end of snout, 6.5
mm.; frontal length (4 mm.) equal to its distance from tip of snout;
anterior chinshields equal to second pair; 4-5 scales touch first
chinshields; labials 4 and 5 enter eye; diameter of eye, 2.4 nun.;
distance between eye and nostril, 3 mm.; scale formula, 26-22-21-
21-18-16. Upper labials, 8-8; two preoculars; nasal divided, the
posterior part very large; ventrals, 173; subcaudals, 38. Total
length, 420 mm.; tail, 54 mm.; length of head, 18 nun.; width of
head, 10.4 mm.

I am indebted to Dr. Leonhard Stejneger and Dr. Doris Cochran
for the privilege of studying these forms in the National Museum.

Boulenger's specimens from Ventanas, Durango and Presidio,
Mazatlan, Sinaloa, appear to agree in general with my specimens.
These three specimens are males with the following combined
(ventral-subcaudal) scale counts: 222, 219, 221; the Guerrero
(female) specimens, 200, 205. This maximum-minimum variation
14-22 is comparable to the variation of 12 in the types (presumably
male and female). The probability is that 7-7 is the normal
formula for upper labials in western Mexican specimens.

374 The University Science Bulletin

Hypsiglena torquata dunklei subsp. nov.

(Plate XXXVII, fig. 1)

Holotijpe. :\ICZ. No. 42594; collected August 10, 1934, Haci-
enda, La Clementina, near Forlon, Tamaulipas, by David Dunkle.

Diagnosis. The most northern variant of Hypsiglena torquata,
varying in the following characters from the typical form: Rostral
bent far back over the snout, which is somewhat compressed (wedge-
like I rather than rounded; the length of the part visible above more
than three fourths its distance from the frontal; prefrontals subtri-
angular, rather than square, due to the fact that the rostral enters
between them, reducing the length of the suture between them; pos-
terior chinshields largest; lower labials, 11-11; loreal irregularly
shaped, not square; ventrals, 162, subcaudals, 57; males with tuber-
cular knobs on scales on side above anus; vertical diameter of eye
minutely less than one half interorbital distance.

Description of the type. Head rather depressed, the snout ex-
tending beyond mouth 1.3 millimeters; width of rostral (2.6 mm.)
mucli less than total length (3.6 mm.), the upper part reflected back
over the snout, its posterior point wedged in between the internasals;
length of rostral seen above equals three fourths or more of its
distance from frontal; suture between the internasals about one half
of the length of the scale, length of internasal about two thirds the
width (1.5X1 mm.); prefrontals (2.1x1-6 mm.) wider than long;
frontal longer than wide (3.3X2.4 mm.), longer than its distance to
the tip of the snout, and about equal to length of the parietal, which
is equal to the distance from parietal to the internasals; nostril be-
tween two nasals, chiefly in the anterior; loreal irregular, much
longer than high; two i>reoculars, upper very large, widely separated
from the frontal, the lower wedged in between the third and fourth
labial; two postoculars, upjicr largest; temporals, 1+2+3; (1+3+
4); fourth and fifth upper labials enter orbit; mental wider than
long; lower labials, 11-11; five touch the anterior chinshields, which
are somewhat shorter than the posterior; posterior chinshields al-
most wholly separated l)y two scales; six scales between first wide
ventral and last lower labial; scale formula, 30, 21, 21, 19, 17;
ventrals, 162; anal divided; subcaudals, 57. Scales generally smooth,
save that those on sides in anal region have tubercular knobs or
keels (probably present only in males) ; a single apical pit present.
Color (in alcoliol). Ground color light buff, with a series of
dorsal spots of brown, the spots, separated by narrow light lines
about a scale wide, are about the length of three scales; occasionally

Taylor: IMexican Snakes 375

one half of the spot will tend to alternate with the other half; a
large dark-brown or purjilish-brown spot on the neck, about the
length of seven scales, reaching laterally to outer scale row; this pre-
ceded by a white or cream collar inA'olving the posterior fourth of
the parietals, not interruj^ted on side of neck or medially; snout and
head liecked with brown, less dense on the parietals; a wide brown-
ish band from eye to angle of mouth; brownish flecks on anterior
upper and lower labials; two rows of lateral spots alternating with
the dorsal row; 43 spots on body; 23 on tail; below immaculate.

Measurements (in mm. I. Snout to vent, 314; tail, 59; tail, di-
vided by total length, .154; head length, 12.2; width, 8; length of
eye, 2; eye to nostril, 2.1.

Remarks. There are eight smaller maxillary teeth preceding two
large fanglike teeth; the fangs bear no trace of grooves. The ventral
count is lower, the subcaudal count higher than any specimens I
have seen of torquata torquata.

I am indebted to ]\Ir. A. Loveridge for the i)rivilege of studying
this specimen. I dedicate it to Mr. David Dunkle, the collector.

376 The University Science Bulletin

PLATE XXXV

Fig. 1. Trimorphodon bi^scutatm (Dumeril and Biliron). EHT-HMS, No.
5339. Hda. EI Sabino, Uriiapan, Michoactin. Raymond Bresson, collector.
Total length, 875 mm.

Fig. 2. Trimorphodon upsilon Cope. EHT-HMS. Xo. 4569. Near Mag-
dalena, Jalisco, Elev. 1300 m. H. M. Smith, collector. Total length, 656 mm.

Fig. 3. Trimorphodon paucimaculatu.^ Taylor. Type. EHT-HMS, No.
4570. (Field number 709.) Near Mazatlan. Sinaloa. E. H. Taylor, collector.
Total length, 880 mm.

Fig. 4. Trimorphodon lambda Cope. Topotype. EHT-HMS, No. 4572.
Near Guaynias, Sonera. E. H. Taylor, collector. Total length, 788 mm.

Taylor: Mexican Snakes

377

PLATE XXXV

*■. %

"•-.

•4

v*»^V

378 The University Science Bulletin

PLATE XXXVI

Fig. 1. Trimorphodon vandcnburghi Klauber, K. U. Museum. No. 8497.
Near San Diego, Cal. E. H. Taylor, collector. Total length, 485 mm.

Fig. 2. Trimorphodon latifascia (Peters). EHT-HMS, No. 5439. Twelve
miles south. Puente de Ixtla. Morelos. E. H. Taylor, collector. Total length,
256 mm.

Taylor: Mexican Snakes

379

PLATE XXVI

^^^^. ^^

^^"•V .Jtk^J^ ^

?**"'' XV

-;/:

I.

z.

380 The University Science Bvlletix

PLATE XXXVn

Fig. 1. Hypsif/Iena torquain dunl-Ui subi^p. nov. MCZ, Xo. 42594. Type.
Forlon, Taniuulipa.s. David Diinkle, collector. Total length, 373 iiiiu.

Fig. 2. Hyp^iglena affiuis Boiilenger. EHT-HMS, No. 4601. Near Mag-
dalena, Jalisco. H. M. Smith, collector. Total length, 279 mm.

Fig. 3. Hyp.^iglcua torquata torquata. EHT-HMS, No. 5200. Twelve miles
south of Puente de Ixtla, Morelos. E. H. Taylor, collector. Total length,
460 mm.

Taylor: Mexican Snakes

381

PLATE XXXVn

/

382 The University Science Billetin

PLATE XXX\11I

Trimorphodon vilkinsonii Cope.

Blanchard collection; collected three miles northwest of El Paso, Texas,
June, 1936. (About natural size.)

Taylor: Mexican Snakes

383

PLATE XXXVIII

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 17

New Species of Mexican Tailless Amphibia

EDWARD H. TAYLOR,
Department of Zoology, University of Kansas

Abstract: This paper describes five new species of Mexican aniirans: Hyla
rickardsi, Potrero Viejo, Veracruz (related to H . loquax) ; HyJa arborescandens,
near Acultzingo, Veracruz (related to Hyla miotytnpanum) ; Rana sierrama-
drensis Agua del Obispo, between Rincon and Cajones, Guerrero (related to
R. palmipes and R. zeteki) ; Eleutherodactylus cactorum, 20' miles northwest of
Tehuacan, Puebla (related to E. axigusti) ; Eleutherodactylus natator, Tlilapam,
Veracruz (related to E. gucnthcri and E. riigrdosa).

FIVE species of tailless amphibians are described in this paper
from the collections made in Mexico by Doctor Hobart Muir
Smith and myself. They may be characterized as follows:

Hyla rickcn'dsi sp. nov.

(Plate XLI, figs. 1-8)

Type. EHT-HMS, No. 5947, ^ ; collected near Potrero Viejo,
Veracruz, August 29, 1936, by Mr. and Mrs. Dyfrig McH. Forbes
and E. H. Taylor.

Paratypes. EHT-HMS, Nos. 5897-5970; same locality, date, and
collectors as type; Nos. 1431-1490, four miles east of Encero, Vera-
cruz, July 17, 1932; E. H. Taylor and H. M. Smith, collectors.

Diagnosis. A medium-sized species most closely related to Hyla
loquax, but differing in having a canthus rostralis, in having the
posterior part of femur pigmented (reddish, lacking dark pigment
in H. loquax), in having a white line along outer edge of foot and
across anal region; the eye and tympanum are proportionally larger.

Vomerine teeth small, directly between choanae; diameter of
tympanum about one half length of eye; head wide, the interorbital
width much greater than width of an eyelid; a medial vocal sac;

(385)
25—4141

386 The University Science Btlletin

fingers more than half webbed; toes with membranes reaching the
base of the disks on the outer side; disk and distal phalanx of the
fourth toe free, save for a marginal skin flap; when arm is placed at
right angles to body an axillary, winglike web becomes evident, ex-
tending along side, reaching two thirds of the distance to elbow;
disks on digits moderately large ; heel reaches to a point between eye
and nostril. Color variable, gray to vinaceous.

Description of the holotype. Head rather broad; eyes prominent,
longer than snout (eye 4.2 mm., snout 4 mm.) ; nostril much nearer
tip of snout than eye, the distance between nostrils equal to distance
from eye to nostril; canthus rostralis rather angular, continued
somewhat in advance of nostrils, the lines when projected intersect-
ing at the tip of the snout; lores sloping but slightly, the region not
concave; diameter of tympanum (2 mm.) one half diameter of eye;
tympanum distinct, but covered with pigmented skin, separated
from eye by a distance equal to two thirds diameter of tympanum ;
upper eyelid (3.2 mm.) much less than interorbital distance (4.5
mm.).

Vomerine teeth on two raised areas, which are much closer to-
gether than to choanae, and wholly between the choanae, which are
distinctly larger than a single group of teeth ; tongue cordiform, dis-
tinctly broader than long, emarginate behind, and free for about
one seventh of its length; openings into the vocal sac very elongate
(4.5 mm.).

Digital pads on fingers large, those on three outer fingers equal to
or larger than tympanum; fingers one half to two thirds webbed;
nowhere do the webs reach the disks save by a narrow margin of
skin; distal subarticular tubercles large, all single; proximal tuber-
cles on two outer fingers more or less obsolete; palm and webs
strongly areolate; a large bean-shaped pad at base of first finger;
a pair of small palmar tubercles; no fold or tubercles on under side
of humerus; legs long, moderately slender, the tibiotarsal articula-
tion reaches to nostril; toes with disks smaller than those on outer
fingers, webbed for three fourths of their length, the web reaching
the disk on outer side of first to third toes; a prominent inner and a
low, rather indistinct, outer metatarsal tubercle; undersurface of
toes, sole of foot and web strongly areolate; a distinct tarsal fold;
a strongly defined supratympanic fold extending back to above arm
or farther along side. When femur is extended a well-defined axil-
lary web is seen extending about half the length of body, and reach-
ing much more than halfway to the distal end of the humerus. A
large vocal sac.

Taylor: Mexican Tailless Amphibia 387

Skin above smooth (under a lens, minutely corrugated) ; skin on
chin is greatly folded, but smooth; chest and abdomen, ventral sur-
face of thigh, and region below anus strongly granular or areolate.

Color. Above vinaceous with numerous spots of a lighter shade
and a few scattered black flecks; below yellowish or cream; con-
cealed surfaces of limbs, and axillary web yellowish; pigment on
sides of head slightly darker tiian on back; a white line borders the
outer edge of foot and fifth toe; a transverse white line above anus;
an indistinct light line borders the canthus and the supratympanic
fold. Femur with scattered pigment on dorsal and posterior sur-
faces; the pigment extends about halfway on the anterior surface
of femur; a few scattered, indistinct, whitish or cream spots along
the sides; limbs more or less barred with brown; dark-brown stripe
on underside of heel, encroaching on sole and underside of fifth toe;
remainder of undersurface of foot, and hand without pigment; outer
part of the chin with pigment; vocal sac lemon yellow.

Measurements {in mm.) . Snout to vent, 37 ; snout, 4; head length,
11.2; head width, 13; eye length, 4.2; eyelid width, 3.2; interorbital
width, 4.5; tympanum, 2; foreleg, 23.6; longest finger, 11; hind leg,
66.5; tibia, 21; foot, 27.5.

Variation. Color variation is great, many of the specimens being
a light gray (yellowish gray at night) with the banding on the limbs
obscure; others are lavender grayish, lavender, clay or light varie-
gated brown. The color is not associated with sex. In all cases
there are small black flecks scattered on the back ; the white line on
the foot is always present, and lighter lines usually follow the
canthi and intersect on the snout.

Remarks. This species has been encountered only twice. For-
tunately each time they were found breeding and large series were
obtained. The specimens were taken from plants growing out of
the water. Many were found clasping, but no eggs were seen. The
calls of this species were mingled in a chorus of the voices of Bufo
marimis, Bufo vallice'ps, Hyla staufferi, Fana pipiens, Rana palmipes
and Hylella picta.

Specimens of this form have been compared with the type of
Hyla godmani Giinther by Mr. H. W. Parker,* who points out
several differences. This form, which Kellogg has placed in the
synonymy of H. miotympanum, lacks an axillary web, and the males
have a horny excrescence on the thumb which is likewise true of
H. miotympanum. From Peters' H. microtis, which Kellogg places
in the synonymy of H. miotympanum, it differs in having the

* I wish to acknowledge my debt to Mr. H. W. Parker for this courtesy.

388

The University Science Bulletin

posterior and anterior surface of the femur without pigment and
the line on side absent. Other differences likewise obtain.

The only other Mexican frog with which it might be confused is
Hyla loquax, which likewise has an axillary web, but which differs
in numerous characters, the most salient of which are mentioned in
the diagnosis. H. loquax is known in Mexico only from Tres Brazos
and Encarnacion, Campeche, where Dr. Hobart Smith discovered
it, September 17-19, 1936.

This species is dedicated to Mr. John A. Rickards, Tacubaya,
Distrito Federal, who has manifested interest in my studies in
Mexican herpetology, and who has presented me with numerous
valuable specimens.

Hyla arborescandens sp. nov.

(Fig. 1)

Tijpe. EHT-HMS, No. 3135, ^ ; collected on mountainside about
3 km. southwest of Acultzingo, Veracruz. July 22, 1932, by E. H.
Taylor and H. M. Smith.

Diagnosis. A medium-sized, olive-colored hyla with strongly-
defined canthi which continue onto tip of snout and intersect;

Wk^-%:^.^'m.

mM mi^

'cSiuaiS*;

Fig. 1. Hyla arborescandens sp. nov. Tvpe. X-2
EHT-HMS, No. 3135, 3 km. southwest of Acultzingo,
Veracruz. Actual snout-to-vent length, 37 mm.

tibiotarsal articulation reaches slightly in advance of the eye;

tympanum rather indistinct, its diameter about one third of the eye;
interorbital space one third wider than eyelid; fingers about one
fourth, toes a little more than two thirds webbed; subgular vocal

sac; throat and chin pustular; abdomen and breast strongly areolate;

dorsal surface with minute corrugations.

Taylor: Mexican Tailless Amphibia 389

Description of holotype. Vomerine teeth in two raised clusters
much closer to each other than to the choanae, situated between but
extending a third of their length behind posterior edges of choanae ;
tongue broad, cordiform, nearly two thirds the width of the mouth,
the surface with prominent papillae; head as broad as or a little
broader than body; eyes prominent, extending beyond profile of
jaw when seen from abo\'e; diameter of the eye (4.2 nmi.) equal to
snout, greater than distance of eye from nostril; distance between
nostrils (3 mm.) less than interorbital distance (4 mm.), which is
greater than the width of an eyelid (3.3 mm.) ; tympanum 1.3 mm.
long, 1.4 mm. high, about one third diameter of eye, separated from
eye by a distance (2.3 mm.) much greater than its diameter; snout
(seen from above) terminating in a sharp point; a slight vertical
ridge on tip of snout; loreal region subvertical, not or but slightly
concave.

No trace of an axillary web; arm normal; digits with tips dilated
into disks, all save those on inner finger larger than tympanum;
second and fourth fingers of ecpal length; fingers one fifth to one
fourth webbed, the webs continued as a narrow dermal fringe along
the edges of fingers to disks; a large nuptual callosity, covered with
minute horny spines, dark brown in color, reaches to near the disk
of first finger; elongate pad on first finger nearly covered with the
nuptual spinosities; two small palmar tubercles, one partly anterior
to the other; outer subarticular tubercles large, flat, none divided;
proximal tubercles small; supernumerary tubercles forming in-
definite rows on metacarpals; a more or less continuous row of
tubercles on under surface of forearm; upper surface of hand and
arm with indistinct, but numerous, flat, pustular elevations. When
legs are folded at right angles the heels overlap about four milli-
meters; disks on toes slightly smaller than those on fingers; those
on three outer toes distinctly larger than tympanum; toes between
two thirds and three fourths webbed, the outer webs excised to level
of the median subarticular tubercle on the fourth toe ; web nowhere
reaches the disks save as a narrow deraial fringe ; distal subarticular
tubercles large, proximal ones small; supernumerary tubercles
forming indistinct rows on metatarsals; a large, flattened inner
metatarsal tubercle, a small indistinct outer; a tarsal fold moder-
ately distinct; third finger only minutely longer than fifth. A
moderately thick fold from behind eye above tympanum, curving
down to the arm insertion; anal flap broad, somewhat thickened,
with numerous grooves on its outer surface; throat pustular; breast

390 The University Science Bulletin

and abdominal surface strongly granular; sides indistinctly areolate;
underside of femur with an area of large, areolate granulations
intermingled with smaller granules, these reaching up on posterior
part of femur to anus, where they form a transverse row of large
pustules; a fold across chest.

Color. Above deep purplish lavender, gradually becoming less
heavily pigmented on sides of head, body and limbs; ventral sur-
faces brownish-cream, with a faint scattering of pigment on ab-
domen and chin; ventral surfaces of heel and foot rather heavily
pigmented except on inner toes; sides lightly pigmented, with a few
very indefinite darker spots exposed; dorsal surface of foot with
small, scattered, darker spots. (In life the color was indefinite
olive, the ventral surface yellowish-cream.) Posterior side of femur
lightly pigmented; a few indefinite lighter spots on dorsal surface
of femur, and on sides of anus; no bands on arms or legs.

Measurement {in mm.). Snout to vent, 37; head length, 12;
head width, 13; snout, 4; foreleg, 26; hand, 13; hind leg, 63; tibia,
19.5; foot, 27.

Remarks. The separation of this species from known Mexican
species may be made by use of the character of the canthus rostralis
combined with the large nuptual callosity and the lack of bands on
hind limb. From H. bistincta, which has a large nuptual callosity,
it may be distinguished by the absence of the very peculiar ventral
prolongation of the anal flap; the presence of a well-defined outer
metatarsal tubercle, and a smaller, more posterior series of vomerine
teeth ; from H. plicata by the posterior position of the vomerine teeth
and the presence of a web on fingers (absent in H. plicata) ; from
H. rickardsi, by the absence of an axillary web; from H. taeniopiis.
by the fact that the pads on outer fingers and toes are much larger
than the tympanum; tongue distinctly notched behind, and a trans-
verse fold on chest; hind limbs not barred; from H. miotympanum
by the more posterior position of the vomerine teeth; the presence
of sharp canthi, the corrugated dorsal surface, the absence of the
light external stripe from elbow and heel to digits. There is no
stripe on upper lip or side. {Hyla miotympanum has been taken
in type locality of the present species.)

The type was collected at night during a downpour of rain, in a
low tree, near the large spring which issues from the side of the
mountain near where the highway crosses. The call was a single,
rather low, mournful call, lasting about one and one-half seconds and
repeated at intervals from three to four minutes. The specimen

Taylor: Mexican Tailless Amphibia

391

was located only with the greatest difficulty. It was heard on the
night of July 21, at which time two hours were spent trying in-
effectually to locate the specimen in a low bushy tree. The follow-
ing day the tree was cut down, but the frog was not found. The
following night the animal was again heard, and finally located in
an adjoining tree, and captured. Specimens of what I believed to
be this species were heard about September 1, 1936, in the forest
trees on this same mountain side, but none was captured.

Eleutherodactylus cactorum sp. nov.

(Fig. 2)

Type. EHT-HMS, No. 6383, 2 ; collected, km. 226, 20 miles
northwest of Tehuacan, Puebia, August 30, 1936; E. H. Taylor,
collector.

Paratypes. EHT-HMS, Nos. 6376 (topotype), H. Radclyffe
Roberts, September, 1936; 6382 (topotype), E. H. Taylor.

(Fig. 2)

Fig. 2. Eleutherodactylus cactorum .sp. nov. Type. EHT, No. 6383, $ , 20
miles northwest of Tehuacan, Puebia. Snout to vent, 77 mm.

392 The University Science Bulletin

Diagnosis. A member of the group in the genus Eleutherodac-
tylus having club-shaped digits with the terminal disk lacking; the
dorsal and ventral surfaces of the terminal phalanx undifferen-
tiated from skin on remainder of digit, and lacking a trace of a
transverse terminal groove; a well-defined ventral disk on abdomen.

A strong fold from eye above tympanum, continuing back to groin
and anus; a transverse postorbital fold; head much broader than
long; eye a little longer than its distance from nostril; upper eyelid
much wider than interorbital space; tibiotarsai articulation reaches
tympanum; hind legs folded at right angles to the body, the heels
are separated by 7.3 mm.; no trace of a tarsal fold; vomerine teeth
between and behind choanae; first finger distinctly longer than sec-
ond. Brownish-white with a pattern of dark lavender on dorsal
surfaces; unifomily cream or yellowish on ventral surfaces.

Description of the holotype. Head much broader than long (31.5
mm.X24.9 mm. I ; interorbital distance distinctly less than upper
eyelid (5 mm. to 8 mm.) ; length of eye (9 mm.) a little greater than
distance from nostrils, which are situated very near extremity of
snout; distance between nostrils less than their distance from eye;
tympanum, 5.8 mm. high, 5 mm. long, separated from eye by a dis-
tance equal to about one third its heiglit ; tympanum partly concealed
by the supra- and posttympanic folds ; no canthus rostralis, the lores
slightly concave, sloping obliquely to lip edge; eyes not extending
beyond outline of head seen from above; tongue broad, not or but
slightly nicked behind; vomerine teeth in a transverse series on
raised, more or less rounded areas, situated between and partially
behind choanae, separated from each other by a distance less than
half their distance from choanae. (Male with a median vocal sac.)
Limbs rather short, the first toe longer and much thicker than second
toe, nearly as long as the third toe; subarticular tubercles on hand
prominent, rounded; a large tubercle at base of first finger and a
large semidivided palmar tubercle ; several supernumerary tubercles
on anterior part of palm; no trace of a web on hand and no lateral
fringes on fingers, save a slight fold on the inner side of digits two
and three; terminal phalanx of digits thickened, not or only slightly
dilated; hind limbs relatively short, thick, the heels separated when
adpressed; digits relatively short, the terminal phalanx not wider
than digit, but thickened somewhat; third toe a little longer than
fifth, which is free for a length half the free part of fourth toe; no
trace of web or dennal folds on digits ; tip of toes thickened, but not
widened; subarticular tubercles prominent; several supernumerary
tubercles on the sole, and occasional low ones between subarticular

Taylor: Mexican Tailless Amphibia 393

tubercles; inner metatarsal tubercle large, about four fifths the
length of first toe; outer metatarsal tubercle somewhat smaller than
inner; a faint trace of a medial metatarsal tubercle; a small super-
numerary tubercle lies between the anterior edges of the metatarsal
tubercles.

Skin above everywhere finely and evenly corrugated, without tu-
bercles; a strong glandular fold begins at posterior corner of the eye,
passes diagonally across upper posterior edge of tympanum and is
continued as a narrow skin fold to anus; a small post tympanic fold
to angle of jaw^; sides and ventral surfaces perfectly smooth, save
for heavy granulation of the median ventral and posterior part of
femur, the granulation reaching up on both sides of the median
posterior groove to the anus, where the granulations become pus-
tular; ventral disk moderately distinct.

Color in life. Ground color gray to brownish-cream with an irreg-
ular pattern of large purplish to lavender blotches, less distinct on
sides; arms and legs banded with dirty cream and lavender; fingers
and palm of hand cream; ventral surfaces cream with slight pig-
mentation on outer edges of jaws; heel and sole dim lavender; toes
generally cream; posterior part of femur lavender with some darker
mottling.

Remarks. The three specimens of this species were collected in
large mounds formed by a species of cactus. Often the mounds
would have a circumference of thirty feet, a height of two or three
feet, and would contain many hundreds of the thick plants. Usually
the plants or stems grow so closely together that one can scarcely
find an opening large enough to insert a pencil. Occasionally
rodents burrow under or gnaw a passageway beneath the mound.
In this way the amphibians apparently gain entrance. When first
picked up the largest female specimen uttered a curious sc^ueal or
scream (suggestive of the noise made by a frightened rabbit), and
voided urine in a continuous stream. The other female behaved in
much the same fashion. More than a total of twenty of these large
plants were destroyed, but only two yielded specimens of the new
form. Numerous specimens of insects, snakes, lizards and other
frogs were taken in this same habitat.

Key to the L.\ticeps Group of Eleutherodactylus

-\. Toes with vestige of a web; head wider than body; a sharp-edged tarsal fold.

Yucatan latlceps Dumeril and Bibron,

AA. Toes without vestige of a web ; head not wider than body ; no tarsal fold.

B. Limbs short, thick; eyelid greater than interorbital distance; tibiotarsal
articulation reaches anterior edge of tympanum ; when leg is folded, the heels
are widely separated. Puebla ; 78 mm cactorum sp. nov.,

394

The University Science Bulletin

BB. Limbs longer; tibiotarsal articulation reaches the middle of eye or slightly
farther; when limbs are folded the heels touch or overlap slightly; eyelid
less than interorbital space.
C. Tympanum % to % diameter of eye; dorsal surface smooth; Texas;

90 mm latrans Cope,

CC. Tympanum scarcely more than one half diameter of eye; Guanajuato;
75 mm augusti Duges.

Measurements of Eleutherodaciylns cactorum E. latrans and

E. augusti (in mm.)

Species

cactorum.

cactorum.

cactorum.

latrans.

augusti.

Number

6382

6376

6383

17755*

6378

Sex

cf

9

9

9

9

Snout to vent

67

78

77

77

75

Snout ...

8.2

7.7

9
8.3

9.3
9

11.5
7.5

11.2

Eve lenffth

8.8

7.4

7.3

8

9

8.1

Interorbital width

5.5

7

6
7.3

5

8

7.4
6.1

7

Upper eyelid

8

Tympanum height and width

5.8, 5.1

6.6, 5.3

5.8, 5

6.8, 5.8

5.4, 5

22

24.8

24.9

27.3

24.2

Head width

29

39

11

11.8
8.5

30.2
44

11

8
12

8

31.5
44

11

7.1
12.3

9

30

52.4

12
11
15

12.7

31

47.5

Finger 1

10.3

Finger 2 . .

8.3

Finger 3

14

Finger 4

9.2

Leg

91

93.5

92

103

109

Tibia

30

40

18

4.6
7.2

11

15.6
6

31

40

18

4.8

7.2

10.5

15.7

7

31
42
17.2

5

8

10.8
15.3

6.2

33.5

47.2

19

6.8
9.4

12.5
19
8

32

Foot . . .

46

21

Toe 1

5.8

Toe 2

9

Toe 3

12.9

18.2

Toe 5

8

* Kan.-^as University Museum, Birds and Mammals.

Eleiithcrodnctylus nafator sp. nov.

(Plate XXXIX. fipr. 2: PI. XL)

Type. EHT, No. 6373, $; Tlilapam, Veracruz, August 16, 1936;
E. H. Taylor, collector.

Paratypes. EHT, Nos. 6371, 6372, 6374, 6375. Same data as type.

Diagnosis. A member of the liohyla group, having the toes
partially webbed, with the tips of digits dilated into moderately
large disks; both inner and outer metatarsal tubercles present, a
tarsal fold running little more than half way to heel; eye about

Taylor: Mexican Tailless Amphibia 395

equal to its distance from snout; canthus rostralis distinct; nostril
a little (about %) nearer median tip of snout than eye; interorbital
distance less than an eyelid; first finger equal to or minutely longer
than second; bluish or purplish black on posterior surface of femur,
enclosing yellowish (or cream) spots and reticulations; tibiotarsal
articulation reaches beyond tip of snout; vomerine teeth between
inner edges, but completely behind choanae; a fold from eye to
shoulder; a diagonal lateral fold beginning much behind tympanum
runs half the length of body; a second lateral fold may also be
present; heels overlap when adpressed. Large; maximum size
known, 93 mm.

Description of the holotype. Head oval, with well-defined canthi,
the lines of which, if projected, would intersect at tip of the snout;
the canthal line is slightly interrupted at nostril, and the canthi are
separated by a somewhat greater distance immediately in front of
nostril than behind nostrils; lores shallowly concave, the sides
sloping rather sharply above, then sloping more obliquely down to
lip. Eyes large, projecting somewhat beyond outline of jaw (as
seen from above) ; eye (10.2 mm.) very slightly longer than its dis-
tance to nostril ; latter a half closer to tip of snout than to the eye ;
distance between nostrils equals interorbital distance; width of
upper eyelid (8 mm.) greater than interorbital distance (7.2 mm.) ;
tympanum height (6 mm.) greater than length (5.1 mm.), somewhat
irregularly pearshaped, separated from eye by a distance of four
millimeters.

Tongue large, cordiform, slightly emarginate behind; vomerine
teeth in diagonal series, on a pair of triangular elevations, separated
from each other posteriorly by a distance greater than that between
choanae and the anterior part of the elevation; inner edges of
choanae separated by a distance a fourth or fifth greater than dis-
tance between nostrils.

Foreleg moderately developed, with the fingers terminating in
well-developed disks, much wider than the narrowest part of outer
phalanx (2 mm. to 3.2 mm.) ; the merest vestige of a web between
the first and second fingers, which continues as a narrow skinfold
on the inner part of the second finger; between the second and third
fingers even a smaller trace of a web, but a skinfold is evident on
the inner edge of the third finger. A prominent tubercle at base of
first finger; a large, single palmar tubercle, bifid anteriorly; meta-
carpal tubercles nearly obsolete (evident in young specimens) ; sub-
articular tubercles large, rounded; hind legs rather heavy, heels
strongly overlapping (8 mm.) ; toes with well-developed terminal

396 The University Science Bulletin

pads which are greater in width than toe behind them (3.8 mm;
2.4 mm.) ; outer metatarsal tubercle small, rounded, opposite middle
of inner tubercle, which is rather elongate (5 mm. X 2 mm.) , sliglitly
less than half the length of first toe (9.5 mm.) ; web between toes
extends to upper edge of subarticular tubercle on the first toe; to
the middle of the tubercle on outer edge of second and third toes;
to posterior edge of proximal tubercle on outer edge of fourth,
which is considerably in advance of this tubercle on fifth toe; a
narrow dermal fringe on edges of all toes to disks; on outer side of
fifth toe the fringe extends from a point some distance behind the
proximal tubercle; a strong tarsal fold slightly more than half
length of tarsus. Tibiotarsal articulation brought forward reaches
beyond snout.

Skin rather smooth above, with some slight rugosities on the
rump ; an indistinct fold from eye back to shoulder ; a fold from be-
hind tympanum continues back about half way (or a little more) on
side; an indistinct, broken, dorsolateral fold arises from the heavy
supratympanic fold and continues back less than half the length of
body. Sides of body with scattered flattened tubercles, almost
granular from axilla, to a point half way back on body; chin with
very indistinct granules; a slight fold on breast, representing an-
terior edge of the abdominal disk which is more or less distinct.

Color in life. Above, metallic, bronzy-green with a brownish-
yellow, narrow^ median line from snout to vent; on sides decidedly
reddish in certain areas; groin yellow; upper surface of limbs same
as ground color of back, save slightly more brownish, with some
dull red on feet; chin, lavender- whitish ; sides of abdomen pinkish;
abdomen and ventral (and to some extent the anterior) part of
femur washed with salmon; under surface of feet purplish; side of
head purplish with a dark line on snout along the lower edge of
canthus and supratympanic fold, widening and partially covering
the tympanum, which is brownish; lip lighter, with dark purplish
spots; arm and foot indistinctly banded. A few deep black spots
on shoulder and in groin; femur and tibia dimly barred; posterior
part of femur black, enclosing yellowish reticulations and spots;
ventral surface of chin, breast, femur, and tibia with brownish
mottling or spots.

Measurements {in mm.). Nos. 6373 $ , 6371 5 , 6372 ^^ , respec-
tively: Snout to vent, 93, 71.5, 42; head length, 35, 31," 18.2; head
width, 35, 30.2, 16; eye, 10.2, 9.4, 5.6; eyelid, 8, 6.3, 5; interorbital
width, 7.2, 5.9, 4; snout, 13, 11.3, 6; tympanum, height, length, 6
x5.1, 4.5x4, 4x4.9; foreleg, 42, 38.2, 23.8; longest finger, 15.8, 12.2,

Taylor: ]Mexican Tailless Amphibia 397

8; hind leg, 145, 127, 79; tibia, 48.5, 43, 28.2; foot, 66, 50, 35.5; long-
est toe, 28, 22, 14.

Variation. The tympanum of the male is far larger in proportion
than in the female, though proportionally higher in the female.

The coloration of No. 6371 is practically identical with the type,
save that the belly was yellowish rather than salmon in life, and the
region below the tympanum reddish; No. 3672 was metalic bronze
above, somewhat variegated with dull reddish and greenish, with the
belly whitish. No. 6374, dark olive above, blackish between the eyes
and on an area behind head; tubercle near ear, orange; the limbs
bronze, barred with black; black and yellow on posterior side of
femur.

Remarks. Save for the fact that both Dr. E. R. Dunn and Dr.
Remington Kellogg* pronounced the type of Hylodes Berkenbuschii
Petersf identical to Liyla. rugulosa Cope|, I would have some slight
doubt as to the distinctness of this form from Peters' species, since
they agree in numerous characters. The characters which seem to
separate the species are the following: The nostril is not twice as far
from eye as the tip of snout (about 8 mm. to eye; 6.2 mm. to tip
of snout) ; the first finger is equal or a little longer than second (not
a little shorter than second) ; belly nearly uniformly yellowish or
salmon, lacking dark spots; in No. 6372, a specimen of comparable
length (42, [43] ), the hind leg is about one ninth longer.

From E. rugulosus (Cope) it differs in having a sharp canthus in-
stead of no, or a very rounded, canthus; snout rather sharply, rather
than bluntly rounded; a relatively longer and slenderer body and
longer limbs, with the tibiotarsal articulation reaching farther; the
heels barely touching, and the webbing on toes distinctly less exten-
sive. I believe it is a very much larger species. The largest au-
thentic E. rugulosus seen measures 58.5 mm. head-body length. Kel-
logg's {loc. cit.) specimen, USNM No. 16567 Orizaba, is certainly a
specimen of the species here described, and probably also 71159. I
suspect that the large specimen, Field Museum No. 1482 (65 mm.)
from Achotal, Veracruz, listed by Kellogg (loc. cit.), also belongs to
this form.

Rana sierramadrensis sp. nov.

(Plate XXXIX, fig. 1)

Type. EHT-HMS, No. 3963B; collected near Agua del Obispo,
between Rincon and Cajones, Guerrero, July 1, 1932, E. H. Taylor.

* Kellogg, U. S. Nat. Mus. Bull., No. 160, 1932, p. llti.
t Peters, Monatsb. Akad. Wiss. Berlin, Dec, 1869, p. 879.
t Cope, Proc. Amer. Philos. Soc, XI, No. 82, p. 160.

398 The University Science Bulletin

Paratijpes. EHT-HMS, Nos. 3963A, topotype; 6565, about 9
km. southwest of Mazatlan, Guerrero (km. 337), July 21, 1936;
6566, near Agua del Obispo (km. 350-351), July 24, 1936; 6567, 6568,
Agua del Obispo, August 1, 1936.

Diagnosis. A member of the group of the genus that includes
Rana palmipes and Rana zeteki, but differing from these in having a
very long hind leg, the tibiotarsal articulation reaching much beyond
the snout. The toes are practically fully webbed, bearing well-
developed terminal disks on the toes; dorsolateral line continuous;
canthus rostralis distinct; outer metatarsal tubercle wanting.

Description of the holotype. Head generally oval, almost triangu-
lar; snout projecting considerably beyond mouth; tongue large,
nearly one half total width of head, deeply notched behind and form-
ing two rounded projections behind; vomerine teeth in two raised
diagonal series, each a little larger than a choana and lying between,
but almost wholly behind, the posterior borders of the choanae;
maxillary teeth small, weak.

Head a little longer than wide (34.2 mm. X 31 mm. ) , the snout (12
mm.) as long as eye; nostril equidistant from eye and median tip of
snout, the slight skinfold behind nostril terminating in a rounded
bead posteriorly; nostril situated chiefly below canthus, but its
upper edge tending to interrupt the continuity of the line; canthi
projected forward intersect some distance anterior to the tip of the
snout; width of upper eyelid (7.7 mm.) greater than interorbital dis-
tance (6 mm.) ; tympanum (6 mm.) separated from eye by a dis-
tance of 5 mm.

A well-defined dorsolateral fold continuous to groin ; a fold, arising
on the dorsolateral fold, going diagonally behind tympanum to arm
insertion, more or less distinct; a short glandular fold behind jaw
angle; skin more or less smooth, with a few minute, pearl-like
tubercles scattered on rump (more pronounced in young) .

Forelimbs moderately large, the first finger distinctly longer than
second but about same length as third ; the terminal disk on fingers
poorly developed, scarcely wider than the digits; three large pads
on the palm, all elongate, separate; subarticular tubercles promi-
nent, that on first finger longest; a single small tubercle on each
metacarpal. Hind leg very long, the tibiotarsal articulation reach-
ing some distance (10 mm.) beyond tip of snout; an indistinct fold
from the inner metatarsal tubercle back to heel ; no outer metatarsal
tubercle, the inner moderate, oval, a little more than one third
length of the first toe; webs continued to base of disks, on all toes,

Taylor: Mexican Tailless Amphibia 399

but somewhat excised between them; terminal disks distinctly wider
than digits; subarticular tubercles elongate, well developed; tibia
with some traces of minute glandular folds; part of posterior sur-
face of femur strongly granulate.

Color. Above brown, becoming grayish brown on the sides, save
in head region; a few indistinct darker brown spots scattered on the
dorsal surface; head with an indistinct dark line from tip of snout,
through eye along outer edge of the dorsolateral glandular fold;
from tip of snout diagonally across upper lip to arm, a golden cream
line; a few dark markings on arm suggesting transverse bands; a
dark spot under forearm, and a heavy, dark lavender band on
posterior side of humerus; a series of darker blotches along the
ventrolateral region, bordering the lateral gray region; femur and
tibia strongly banded, with scattered spots on anterior face of both,
the bands on tibia reaching half way across under surface ; posterior
surface of femur black, with yellowish reticulations or spots; foot
with transverse bands; under surface of foot dark purplish.

Measurements {in iiwi.). Nos. 3963B, 3963A, 6565, respectively.
Snout to vent, 91, 70, 44.5; head length, 34.2, 28.5, 17.6; head width,
31, 25.2, 14; snout, 12, 10, 7.5; eye, 11.8, 8.1, 6; interorbital width,
6, 5.5, 3; tympanum, 6, 4.2, 3.1; forelimb, 50, 38, 27; hind leg, 168,
132, 69; tibia, 55, 44.5, 25.5; foot, 72, 55, 31.

Variation. The younger specimens 6665-6668 are reddish bronze
above; posterior part of the dorsolateral line whitish; a broad black
band from tip of snout, involving whole tympanum, continues along
the upper part of side to groin; lower on side, grayish to gray-white.
The line on lip especially distinct, bordered below by brown to
black-brown; on abdomen and throat, cream or dirty white. The
eye of the type is slightly larger than in the second specimen 3963A.
This latter specimen in life had a few small olive spots on the back
which are evident in the picture (plate XXXIX, fig. 1).

Remarks. This large species replaces Rana palmipes in the higher
parts of the Sierra Madre del Sur. It is amazingly alert and makes
prodigious leaps when pursued.

I have had large series of Rana palmipes for comparison from
Veracruz and Chiapas. This species differs in having shorter legs;
the tibiotarsal articulation reaching slightly beyond eye; the heels
barely touching (very strongly overlapping in R. sierraniadrensis) ,
the skin strongly pustulous on sides and posterior part of back; the
pads on the toes smaller; marked dermal fringes along edges of
middle fingers; and the dorsolateral fold not continuous to groin.

400 The University Science Bulletin

PLATE XXXIX

Fig. 1. Raiio sienomadrensis sp. nov. Paratype: EHT-HMS, No. 3963A,
Agua del Obi.spo, between Rincon and Cajones, Guerrero, Mexico. Snout to
vent, 70 mm.

Fig. 2. Eleutherodactylus natator sp. nov. Paratype EHT-HMS, No. 6371
9, Tililapam, Veracruz. Mexico. Snout to vent, 71.5 mm.

Taylor: Mexican Tailless Amphibia

401

PLATE XXXIX

T

'4'

/.

26—4141

402 The University Science Bulletin

PLATE XL

Eleutherodactylus notator sp. nov. Type. EHT-HMS, No. 6373 $ , Tlilii-
pam, Veracruz, Mexico. Snout to vent, 93 mm.

Taylor: INIexican Tailless Amphibl\

403

PLATE XL

/ : . •

-' •^-•■^^'

/

' \'^ ^%

404 The University Science Bulletin

PLATE XLI

Hyla rickardsi sp. nov. (about natural size).

Fig. 1. EHT-HMS. No. 1472. Paratype, near Enceio, Veracruz.

Fig. 2. EHT-HMS, No. 1474. Paratype, near Encero, Veracruz.

Fig. 3. EHT-HMS, No. 5947. Tyi)e, Potrero Viejo, Veracruz.

Fig. 4. EHT-HMS, No. 5942. Paratype, Viejo, Veracruz.

Fig. 5. EHT-HMS. No. 1444. Paratype, near Encero, Veracruz.

Fig. 6. EHT-HMS, No. 14S9. Paratype, near Encero, Veracruz.

Fig. 7. EHT-HMS. No. 5968. Paratype, Potrero Viejo, Veracruz.

Fig. 8. EHT-HMS. No. 5946. Paratype, Potrero Viejo, Veracruz.

Taylor: IMexican Tailless Amphibia

405

PLATE XLI

4- ^-

^

^ .

S

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV | June 1. 1938 [No. 18

A New Aiuiran Amphibian from the Pliocene of Kansas

EDWARD H. TAYLOR,

Department of Zoilhigy, linivprsity of Kansas

(Plates XLII-XLV)

Abstiuct: Scnphiopu-s stiidcri sp. nov. is described from the Middle Pliocene
of Lojian county, Kansas. It is comiiared in detail with Scaphiopus bombijrons
Cope, a modern species of the genus.

DrRIXCi the Slimmer of 193G a lu.^.-^il frog ^vas presented to Mr.
Claude Hibbard for the Museum of the University of Kansas
by Mr. Frank Studer. The specimen had been obtained by Mr.
Studer some time previously from a bed of (hatomaceous marl lo-
cated in Logan county (sec. 7, T. 11 S., R. 37 W.) about a mile east
of the "Rliino Hill Quarry," which is near the east line of Wallace
county.

This bed, some feet in thickness, of a gray-white color, has yielded
a large number of fossil fresh-water fish, but so far as I know this
is the first and only representative of any other vertebrate group
that has been found.

The amphibian was obtained by clea\'ing a slab of marl. In doing
this, portions of the skeleton remained on both sides of the slab.
When first obtained the fossil was perfect, but its subsecjuent treat-
luent has brought about the loss of certain of the bones, including
important skull elements and the greater portion of the vertebrae.
However, many of the bones are still intact and the bone impressions
are relatively clear, save in the skull region.

The specimen is certainly referable to the family Scaphiopodidae,
and because of its resemblance to certain modern forms I am pro-
posing tentatively to place it in the genus Scaphiopus.

(407)

408 The University Science Bulletin

Scaphiopiis studeri sp. nov.

Holotijpe. Univ. Kansas Museum of Paleontology No. 1478, con-
sisting of a split slab of marl each portion of which contains both
imprints and bones of a single animal.

Type locality. Logan county, Kansas (sec. 7, T. 11 S., R. 37 W.),
about a mile east of "Rhino Hill Quarry," near the Wallace county

line.

Horizon. Diatomaceous marl, in contact with the Eclson Beds,

]\Iiddle Pliocene.

Diagnosis. A medium-sized Scaphiopiis approaching the character
of Scaphiopus bombifrons Cope more closely than any other living
from; and related to Scaphiopus pliobatrachus Taylor, a fossil form.
Spade large, tibiale and fibulare fused together at ends ; sacral dia-
pophyses fan-shaped, the sacral vertebra solidly fused to the uro-
style; apparently a broad fontanelle between the frontoparietals.
The ethmoid has a rounded knoblike termination. Total length,
snout to vent, about 58 mm.; width of knees with limbs at right
angles to body, 47 mm.

Description of the Holotype. [Photographs of the fossil remains
are given (plates XLII and XLIII) and this is interpreted by a
series of drawings of the individual elements (plates XLIV and
XLV). I have compared these with the same elements in Scaphiopus
bombijrons now occurring in Kansas. Some of the points of differ-
ence are worthy of note: 1st vertebra very different in size and gen-
eral character; carpalia very different; tibiale and fibulare solidly
fused (not so in bombijrons) ; distal ethmoid termination not bending
down, but lying in general plane of the remainder of the bone.
Since the various elements of the fossil and recent forms are figured
side by side, other numerous differences may be discerned, and need
not be discussed here.]

Skull

The skull is in an extremely fragmentary condition. The ventral
part of the braincase is l)roken away, leaving the frontoparietal ele-
ments showing with a large fontanelle between them. Anteriorly a
part of the ethmoid remains. It is somewhat boxlike, its anterior part
enclosed, presenting at its anterior end a knoblike termination. In
the character of the frontoparietals and the condition of the ethmoid
this species resembles Scaphiopus bombifrons. In the latter the
anterior j^yrojection is deflected, while in the present specimen it is
straight. The nasals, turbinals, and vomers are badly shattered and
the few remaining fragments tell little of the former extent of the

Taylor: A New Axuran Amphibian 409

bones, and their relationships. A triradial fragment of tlie pterygoid
is present, in contact with tlie niaxilhiry.

No traces of vomerine teeth are discernible on the one or two
fragments \\hich may represent portions of the vomers. (Plate
XLIV, fig. 2.)

Maxillary. Portions of the maxillaries are present on both sides.
On the left side is a fragment measuring 5.2 mm. along the labial
edge; it contains 15 teeth with one or two missing tooth spaces; on
the right side the maxillary consists of four fragments which lie in
a straight line (rather than curving) and show^ 39 teeth or tooth
spaces with a total length of 9.1 mm. The maxillary shelf is at
right angles to the direction of the teeth with a very slight trough
standing out about six eightlis mm. from the bone. The impression
of the posterior extension of the maxillary shows the total length to
be 10 mm., the posterior part of the bone extending out from the
shaft of the quadrato-jugal to which it attaches; the greatest width
(vertical) of the maxillary (2.(i mm.) is near its most anterior point.
(Plate XLIV, figs. 5 and 6.)

Premaxillary. A portion of one premaxillary is present, but it has
been displaced backward out of line with the maxillaries. It bears
8 teeth and two spaces. The process is missing.

Frontopaiietals. These are thickened elements with a clearly de-
fined longitudinal fontanelle between them suggesting the condition
obtaining in Scaphiopus bomblfrons. Their depth in the matrix
shows a skull with considerable depth as is typical of modern
Scaphiopus.

Parasphenoid. This element is completely shattered and only one
or two small fragments are present.

Ethmoid. This element is represented by its anterior end, which
is boxlike, presenting anteriorly a knoblike boss cjuite unlike the
proboscislike termination of this element in S. bombifrons. Other
skull elements cannot be made out in their entirety, although I be-
lieve all were present when the specimen was first obtained, with the
possible exception of the mandible. (Plate XLIV, figs. 12, 13.)

The Vertebral Column

The vertebral column, having a total length to end of the urostyle
of 35.2 mm., consists of 9 vertebrae, with the urostyle solidly fused
to the ninth (sacral) vertebra. The diapophyses are strongly di-
lated, fan shaped. The vertebrae from the second to the ninth are
so shattered that it is difficult to interpret all the characters from
the fragments and the impressions.

410 The University Science Bulletin

It is impossible to determine, beyond peradventurc, the characters
of the centra; however, the impression left of the ventral surfaces
of the vertebrae show the suture between them curved so as to
suggest strongly that most of the vertebrae are procoelous.

The first vertebra is complete, and seen from below shows two
curving emarginations on the anterior upper edge of the vertebra
with a V-shaped median notch. The anterior sockets for the condyles
arc large, and the element is strongly widened anteriorly and
narrowed suddenly posteriorly. This vertebra is opisthocoelous.

The transverse process of the second vertebra is directed slightly
forward; it is not or but sligiitly widened. The third vertebra has
a transverse process distinctly wider distally than proximally,
strongly compressed vertically at its proximal end; on the fourth
vertebra the process is wider at the base than distally, terminating
in a more or less rounded tip directed almost directly outward; its
length is 4.3 mm. The fifth and sixth processes are very short, 1.2-
1.4 mm. in length.

The sacral vertebra has the ventral part of the centrum shattered
and missing; however, most of the urostyle is present and the greatly
widened processes of the sacral vertebra. These elements are fused
together. The greatest transverse width of the sacral vertebra is 10
mm.; the greatest width of the transverse process 5 mm.; the length
of vertebra and urostyle (estimated) 16.3 nun. Tlic urostyle is
compressed vertically and is rounded below; it is without any ventral
crest. (Plate XLIV, fig. 4.1

The Pectoral Girdle

The pectoral girdle is represented by a few fragments and im-
pressions. The coracoid is strongly bowed ventrally and at the same
time tends to curve anteriorly ; the clavicle is slender and a nearly
complete impression is discernible. The median point of contact of
the clavicles is at their extreme tips, and there is no suggestion of an
overlapping. In the case of the coracoids I can make out the median
end of neither. There is no evidence of a cartilage or bone posterior
to the coracoid (omosternum), but there are some bone fragments
just anterior which I cannot certainly interpret. (Plate XLV, fig. 12. )

Scapula. This is discernible only as a deep depression which
shows the element narrowing typically near the middle of the bone,
then curving out strongly. A proximal portion of the suprascapula
is present and the outline of the complete bone is shown by the im-
pression left l)y the remaining part. The proximal half of the ele-
ment is rectangular, but the distal part narrow and the anterior edge

Taylor: A Xkw Aniran Amphibian

411

of the complete element is distinctly curved, r;ither liki' the con-
dition in modern Scaphiopus Iwmbifrons.

Arm

Humerus. The humeri are more or less broken, but where broken
the imprint is clear enoujih to show the following measurements:
Total length, 16 mm. and 15.8 nun.; greatest width, 4.6 nmi. and
4.8 mm.; diameter of the ball at distal end, 2.3 mm.; each of these
elements has a compressed ridge or crest on its anterior half, whicli
extends nearly two thirds the length of the bone. The diameter of
the proximal head of the humerus is 3 mm. (Plate XLV, fig. 13.)

Radio-idnd. This element is now missing save for a few fragments.
The impression shows a bone 10.5 mm. long (right side), 10.2 mm.
(left side) ; each 3.5 mm. wide at the distal end.

At the distal end of the ulnar part is an epiphysis separately
ossified. I It appears to be the cartilagenous element present in
modern Scaphiopus usually so conspicuous because of its pure white
color.) (Plate XLV, fig. 9.)

Carpalia. For the position of these elements, refer to the draw-
ings. There are eight or nine of these, including the distal elements
of the prepollex. The ulnare of the left hand seen from the dorsal
surface is soinewliat compressed. Its exposed dorsal surface shows a
median groove with two ridges bordering it ; the centrale is more
than double the size of the ulnare, its outer face a curving surface.
On the dorsal surface there is a slight depression, roughly the shape
of an hour glass. The radiale is apparently missing save for a small
portion of the element which presents a rounded convex surface
which fits into a similar socket in the preceding carpal element.
Apparently a small carpal is lost from the proximal end of the first
finger; below this apparently missing element is a very tiny element
imbedded in the matrix. The prepollex bone is 1.8 nnn. long, and
1.15 mm. wide at the base. It narrows rapidly to a rounded ]ioint,
curving slightly. I Plate XLV, figs. 3, 11. )

Measurements of metacarpals and phalanges (right). (Plat(> XLV, Fig. 3.)

First finger. .
Second finger
Third finger .
Fourth finger

Meta-
carpal.

5

4.3
4.9
4

r"irst
phalanx.

0.5
1.5
2.2
1.1

Second
phalanx.

0.5
0.5

1.2
0.85

Third

phalanx.

0.5
0.5

412 The University Science Bulletin

The Pelvic Girdle

Ilium. A portion of the ilium is present on both sides. On the
right side the widened acetabular part is broken just anterior to the
acetabular depression. The total length of this element is 23 milli-
meters to the middle of the acetabulum (estimated from the impres-
sion). At the acetabular end the bone widens. There is no knob
or projecting elevation of the dorsal posterior part. The anterior
ends of the ilia reach and fuse with the sacral vertebra, but do not
extend to the anterior edge of the widened processes. (Plate XLIV,
figs. 8, 10.)

Ischia. The right and left ischia are closely fused, and where they
join with the ilium they form an elevation, when these elements are
seen in profile.

Pubis. This element seems to have been at least partially of soft
cartilage, for there is a discontinuity of the ventral outline of the
pelvis in lateral view; no trace of a suture can be discerned to sug-
gest fusion with ischium. It may lun'e been normally missing or
not ossified. The edges of the acetabular cup are raised on both the
ischium and ilium.

Hind limb. Both hind limbs are present and all of the bones are
likewise present on one or the other. The larger, thicker parts are
more or less shattered. On both sides the feet are folded so as to
show their ventral surfaces.

Femur. The left femur has the acetabular head complete. The
greatest diameter of the head is about 2.2 mm.; the total length of
the femur is 21 mm., and its greatest diameter 4.2 nnn. at the distal
end; it narrows to a width of 1.5 mm.; the posterior edge is slightly
flattened, forming a crest (this portion is broken, but an impression
of the crest can be discerned). Near the distal end is a thickened
boss. (Plate XLV, fig. 2.)

Tibia-fibula. This element, somewhat broken at each end, has a
total length of 18.2 mm. The arterial foramina on opposite sides of
the shaft are distinct ; that of the fibula one millimeter in advance of
the other. No very marked grooves are visible between the fused
elements, due to the shattering of portions of the exposed surface.
Tiie internal partition between the two elements is discernible at the
proximal end where the bony surface is broken. (Plate XLV, fig. 10.)

Tibiale and Fibulare (astragulus and calcaneumL These two ele-
ments are present on the left side only. They are apparently com-
pletely fused together and are perfect save that a fragment is frac-
tured from the iimer distal end. It is still in place, however. The

Taylor: A New Anuran Amphibian

413

distal width of the fused Ixnics is (i iniii.; tlic proximal width, 4.0
mm.; near tlie proximal end of the fibulare, on its ventral surface,
is a slight ridge from near the point of contact with the tibiale, to
its outer edge, limiting a somewhat rougliencd triangular area on
the ventrolateral surface. (Plate XLV, fig. 6.)

Tarsalia. Four tarsal elements, including the prehallux, are pres-
ent— ajijiarently representing the complete eciuipment. A small ele-
ment is present at the proximal end of the first digit, and one at the
proximal ends of the second and third equally in contact with both
digits. The prehallux consists of an enlarged basal element from
whicli arises a broad shovel-shaped element which originally was
pressed against the ventral surface of the proximal phalanges of the
first and second digits. Originally intact, this latter element has
been broken, but the impression in the matrix can be seen.

The two small tarsalia (Nos. 1 and 2 I are flattened, biscuit-shaped.
The third may represent a fusion of two elements. The large spade
(prehallux) had an original length of 4.3 mm. and a width (esti-
mated) of about 3 mm. The sharp cutting edge (probably originally
bearing horn) is evident, (Plate XLV, fig. 6.)

Metatarsals, phalanges. These elements for the most part or their
clear impressions are present. Their measurements in millimeters
from the first to fifth digits arc given in a table.

Measurements of metatarsals and phalanges. (Plate XLV, Fig. 6.)

Meta-
tarsal.

First
phalanx.

Second
phalanx.

Third
phalanx.

Right foot :

5.6

8.1

10.1

11.5

9.3

5.6

8.1

10.1

11.5

9.3

1.7
2.3
3.6
5.2
3.6

1.5
2.2
3.8
5,1
3.8

Third toe

1.7

Fifth toe

2

0.6

Left foot:

Third toe ....

1.95

3

2

Fourth toe

Fifth toe

1.2
0.6

The terminal elements vary in shape. That of the fifth toe is
rather jn'ramichd with a slight cylindrical projection (slightly tri-
angular in profile) from the distal end. That on the fourth finger
is more curving, elongated and clawlike. The others are small,
somewhat triangular, cur\-ing downward very slightly.

Remarks. The relationship of this form to Scaphiopus plio-
batrachus Taylor from a nearby locality in the Edson Beds is
discussed in a paper now in press and is not repeated here.

414

The University Science Bulletin

,;iv

PLATE XLII

ScapJtiupas sttideri sp. nov. Type: Lower part of slab showing bones, with
the exception of the feet, and tibia fibula from a dorsal \iew. Liii)ressions
are of the ventral surfaces of bones. Actual "head-bodv" length about 58 mm.

Taylor: A New Anuran Amphibian

415

./• 'j'^iV ft'"''' yf. "^.'^ .,,»*»r-^/Vr*

n.-^iV;*!

;•'■ ;'yXf;^)^«: ^ ''*'^ K

-w ', (i'> ^ i'* ■ -' ■ '; J ■*' - *"♦ I i» ^^B fa

■■' /5 : Iff /Nit ' -^itW V

'* 'f' ■ i S. f

■'A

r

' •< >}

7 ^f

■ 7 (.

PLATE XLUl

Scaphiopus studeri sp. no\'. Type: Upper part of slab showing bones from
their xcntral surfaces, the impressions from the dorsal surface.

416 The University Science Bulletin

PLATE XLIV

Fifi. 1. ScaphiopuK bombifrons. Morton countv, Kansas. Ventral view ot
skull.

Fig. 2. Scnphiopns studcri sy). no\. Type: Ventral view of skull.

Fig. 3. Scaphiopus bombijrons. Vertebral column.

Fig. 4. Scaphiopus studeri sp. nov. Vertebral cohunn. Ventral view.
Length, 35.2 mm.

Figs. 5 and 6. Scaphiopus studeri sp. nov. Groups of teeth much enlarged.

Fig. 7. Scaphiopus bombifrovs. Pelvic girdle.

Fig. 8. Scaphio'pus studeri sp. nov. Pelvic girdle length, 25.2 mm.

Fig. 9. Scaphiopus bombifrons. Pelvic girdle, lateral view, 22.6 mm.

Fig. 10. Scaphiopus studeri sp. nov. Pelvic girdle, lateral view (the posterior
fragment of the pelvis shown in Fig. 8 was destroyed accidentally before the
drawing of Fig. 10 was made); (length, 23 mm.).

Fig. 11. Scaphiopus bombifrons. Front view of skidl, showing the anterior
part of the ethmoid deflected downward almost in contact with the pre-
inaxillary (much enlarged).

Fig. 12. Scaphiopus studeri sp. nov. Portion of ethmoid as it appears in
situ (much enlarged).

Fig. 13. Same; from a front view of its anterior end corresponding to the
position of ethmoid in Fig. 11, enlarged to same scale as Fig. 12.

Fig. 14. Scaphiopus studeri sp. nov. Impression of the radio-ulna. Actual
size, 10.5 mm. long.

Fig. 15. Scaphiopus bombifrons. Radio-ulna, actual size, 10 mm. long.

Taylor: A New Anuran Amphibian

417

PLATE XLIV

27—4141

418 The Uxiversity Science ]3uli.etin

PLATE XLV

Fig. 1. Scaphiopus homhijrons. Femur.

Fig. 2. Scaphiopus studeri sp. nov. Femur (length, 21 mm.).

Fig. 3. ScapJiiopus studeri sp. nov. Hand.

Fig. 4. Scaphiopus homhijrons. Hand.

Fig. 5. Scaphiopus homhijrons. Foot.

Fig. 6. Scaphiopus studeri sp. nov. Foot (length, 28 mm.).

Fig. 7. Scaphiopus hovihijrons. Pectoral girdle from dor.sal view.

Fig. 8. Same ventral view.

Fig. 9. Scaphiopus homhijrons. Tibia-fibula.

Fig. 10. Scaphiopus studeri sp. nov. Tibia-fibula (length, 18.2 mm.).

Fig. 11. Scaphiopus studeri sp. nov. Carpalia (greatly enlarged).

Fig. 12. Scaphiopais studeri sp. noA-. Upper, bone, clavicle; lower, coracoid.

Fig. 13. Scaphiopais studeri .sp. nov. Humerus (actual length, 16 mm.).

Fig. 14. Scaphiopus homhijrons. Humerus.

Taylor: A New Anuran Amphibian

419

PLATE XLV

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 19

Frogs of the. Hyla eximia Group in Mexico, with
Descriptions of Two New Species

EDWARD H. TAYLOR,

Department nf Zoology, University of Kansas

Abstr.act: Six Mexican forms of the genus Hyla are recognized as belong-
ing in the eximia group. These are Hyla eximia Baird; //. euphorbiacea
Ciiinther; H. lajrentzi Mertens and Wolterstorff ; H. regilla Baird and Girard;
H. cdrdenasi sp. nov. ; and H. umghtoram sp. no^'. Cope's species H. gracilipes
is referred to the synonym of H . eximia, while H. smithii, placed in synonymy
of eximia by Kellogg (1932), is regarded as a. good species belonging in a
different group. Hyliola bocourti Mocquard is referred to the synonymy of
Hyla euphorbiacea Giinther.

TN MY endeavor to identify correctly a collection of Mexican
-■- Hylid frogs, I found considerable difficulty in properly allocating
several forms, obviously related to the small Hyla eximia Baird, but
which differed from this species in numerous characters. After a
review of the literature (which appears to be somewhat confused),
and an examination of the types of Hyla eximia and H. gracilipes, I
have arrived at certain conclusions differing from those of certain
former workers.

The literature offered the following names for these forms oc-
curring in Mexico or adjoining territory (forms related so as to war-
rant their association into a species group which is here designated
as the eximia group): Hyla eximia Baird, Hyla euphorbiacea
Giinther, Hyla gracilipes Cope, Hyliola bocourti Mocquard, Hyla
regilla Baird and Girard and Hyla lajrentzi Mertens and Wolter-
storff.

A critical examination of the cotypes of Hyla gracilipes and the
cotypes of H. eximia shows them to belong in what I am forced to
regard as a single species {H. eximia), since I can discern no con-
stant structural characters or character which will serve to separate

(421)

422 The University Science Bulletin

the forms. Unfortunately most of the color characters have long
since disappeared from these specimens.

The description of Hyliola bocourti Mocquard, based on speci-
mens from Upper Vera Paz, Guatemala, agrees with the brief de-
scription of Hyla euphorbiacea to the extent that it is, at least
tentatively, being placed in the synonymy of that species.

The collection made by Dr. Hobart M. Smith and myself in
Mexico contains about 272 specimens of the group. This material
for study has been augmented by an examination of more than 150
specimens in the National Museum, a collection of some fifteen
specimens in Cornell University, and certain specimens in the col-
lection of the University of Michigan, to the authorities and curators
of which institutions my heartiest thanks are given.

This extensive material consists of six species, one of which, Hyla
regilla, occurring in Baja California and western United States, is
not considered in this paper. Three of the species are recognized un-
der the published names Hyla eximia, Hyla euphorbiacea, and
Hyla lafrentzi. Two are regarded as new, and are described under
the names, Hyla wrightorum and Hyla cdrdenasi.

Key to Mexican Species of the Hyla eximia Group

A. Skill granular above or somewhat pustulate. 42 iniu. Baja California and California.

Hyla regilla Baird and Giiard, p. 422
AA. Skin smooth or minutely corrugate.

B. Posterior, and to a lesser extent, anterior region of the femur and groin with
blackish or brownish reticulation enclosing rounded or irregular yellow-cream
spots. 37 mm. Oaxaca and (? south to Guatemala).

Hyla euphorbiacea Gunther, p. 426
BB. Posterior and anterior thigh regions with sparce, equally distributed pigmen-
tation, lacking yellow-cream spots.
C. Til)iotarsal articulation brought forward reaches to near the eye.*

D. Diameter of eye equal or greater than length of snout; snout
bluntly rounded. Uniform greenish without distinct marks on side
of head and body. Eyelid greater than interorbital distance. 39
mm. Puebla Hyla cdrdenasi sp. nov.. p. 43(1

DD. Diameter of eye distinctly less than length of snout ; snout rather
oval; distinct band on head and side, light -edged above; eyelid
less than interorbital width. 35 mm. Southern Mexico from Puebla
and Morclos to Durango and ? San Luis Potosi.

Hyla eximia Baird, p. 423

CC. Til)iotarsal articulation brought forward reaches the nostril or slightly

beyond the tip of snout.

D. Larger form, green with the lateral band on head and side, and

spots on sacral region, edged with silvery white ; the front side of

the tibia blackish, bordered by a white line above, continued on

foot. .50 mm. Veracruz and Morelos.

Hyla lafrentzi Mertens and Wolterstorff, p. 433

* Note. — If specimens are somewhat dried or shrivelled, or the body is distended with
eggs, the relationship of body length to leg length (as shown by bringing the leg forward)
is often difficult to determine. In such cases the tibiotarsal articulation may fail to reach as
far forward as indicated in the key.

Taylor: Frogs of Mexico 423

DD. Somewhat smalliT fonu with a less-distiiift hand fin i'idc of body,
and dorsal spots not edged with white; anterior edge of tibia with
heavy brown spots. lacking a white line; webbing on toes less than
lafrentzi. 42 nun. Chilmahua, Mexico; New Mexico and Arizona,
U. S. A Hi/la ivrightnrtim sp. nov., p. 43i>

Hyla eximia Baivd

(Plate I.XVI. figs. 1-10: Plate XLVII. figs. 3-5)

Hyla eximia Baird, Proc. Acad. Nat. Sci. Phila., 7. 1854. p. <il (type de.scription ; t\-pe
locality. City of Mexico, Distrito Federal, Mex. Major William Rich, collector); and Kept. U.
S. Mex. BoundaPi- Siuvey, 2, Reptiles, 18.59, p. 29, pi. 38, figs. 8-10; Peter.*, Monatsb. Akad.
Berlin, 1869, p. 880 (part.); Brocchi Bull. Soc. Philos.. Paris, (7), I, p. 128 (part.);
Boulenger, Cat. Batr. Sal. British Mus., Ed. II, 1882, p. 378, (part.); Giinther, Biologia
Centrali-Aniericana, Reptilia, Batrachia, Jim?, 1901, pp. 2f)l-2(!2, (part.); Brocchi, Mission
Scientifique au Mexique et dans TAnierique C'entrale, jiart 3. 2d sec. liv. 1, 1881, pp. 32-33,
111. 13. fig'.. 3, 4, 4a (part.); Cope, Bull. U. S. Xat. Mus., No. 32, p. 14 (part.); Nieden,
Das Tierreich, Amna I, Berlin and Leipzig. 1923, p. 24") (part.); Kellogg, Bull. U. S. Nat.
Mus., No. IfiO, 1932, jip. 1.53, l(i4-l(>8 (part.).

Hyla gracilipes Cope, Proc. Acad. Nat. Sci. Phila.. V. 17, Oct.. ISO."), p. 194 (type
description: type locality, "Tableland northeast of the City of Mexico" [Mirador, Vera-
cruz]); and Bull. U. S. Nat. Mus., No. 32, 1887, p. 14; Brocchi, Mission Scientifique
au Me.xique et dans TAmerique Centrale, part 3. 2d sec. liv. 1, 1881, pp. 36-37; Boulenger,
Cat. Batr. Sal. British Mus., Ed. II, 1882, p. 378; Giinther, Biologia Centrali-Aniericana
Reptilia and Batrachia, June, 1901, p. 262; Kellogg, Bull. U. S. Nat. Mus., No. 160, 1932,
pp. 1.53-1.54 and 168-170 (part.).

Tlie identity of Hyla gracilipes Cope has been something of a
puzzle. Boulenger (1882 loc. cit.) recognizes the form as distinct
and merely copies a part of Cope's description, while Giinther
(June, IDOl, loc. cit.) places the species in the synonymy of Hyla
eximia. He comments that the statement in the description, "eyes
little prominent, one half tympanic disk," is probably a lapsus
calami.

Brocchi (loc. cit.) recognizes the species apparently without hav-
ing specimens, and wholly on the authority of Cope et al.

Kellogg {loc. cit.) revives the name and includes the four cotypes.
and nine specimens from the state erf Chihuahua. He places Hyla
lafrentzi Mertens and Wolterstorff in the synonymy of the species.

An examination of the cotypes (USNM, Nos. 15318-15321, which,
according to Cope, were from "Mexican tableland northeast of the
City of Mexico," and according to the USNM Catalogue, ''Mirador,
state of Veracruz, 1863, Carlos Sartorius, collector") shows four
small specimens, so faded that practically no trace of the original
coloration can be discerned, and the markings are very dim. How-
ever, the general characters of skin, body and limbs are not effaced.
When compared with H. eximia from the type locality region I find
only differences so slight that I regard them as negligible. The
great length of the foot mentioned is but a trifle longer in one speci-
men than the normal for H. exi^nia. In all. the tympanum is only

424 The University Science Bulletin

one half (or ks.-~) than the eye, instead of the opposite (an erroneous
statement given in the type description).

Another significant fact is that all are small, measuring from 28
to 33 mm. That these are adult is evidenced by the fact that two
have the ovaries filled with ripe eggs. I am placing H. gracilipes
Cope as a synonym of H. eximia Baird.

While H. eximia shows numerous variations in its wide range, it
appears that these are of lesser import than would warrant even sub-
specific designations, at least as regards the material available to
me. Specimens from Puebla in the south, with the spotting greatly
reduced, compared to the heavily lined and spotted ones from
Nayarit and Durango, seem superficially to be rather different from
each other, but there is no sharp limit to these forms and the transi-
tion seems gradual in various populations.

The most recent treatment of H. eximia is that of Kellogg {loc.
cit.). He includes Hyla euphorbiacea Giinthcr and Hyla sfmthii
Boulenger in the synonymy. I am commenting on the former in this
paper. I have recently [Trans. Kan. Acad. Sci., 39, 1936 (July 2,
1937), pp. 357-359, pi. 2, figs. 1-5] redescribed H. smithii. Be-
yond question it is distinct from Hyla eximia Baird. Specimens of
these forms have been examined by Mr. H. W. Parker of the British
Museum and drawings made of the types of Hyla smithii which show
the characteristic differences between the forms. H. smithii is re-
lated to Hyla underivoodi and Hylella picta, showing the character-
istic type of pigmentation, entire lack of green coloration, larger
finger pads, greater amount of webbing on the toes and feebly de-
veloped vomerine teeth. H. smithii has a distribution coextensive
with H. eximia in the western part of the range, but probably occu-
pies more territory in Guerrero and perhaps less in the eastern part
of the range of H. eximia in Puebla and Veracruz.

The collection made by Dr. H. M. Smith and myself in Mexico
totals some 245 specimens. The following localities are represented :
Nos. 1840-1877, 3 km. northeast, Cholula, Puebla, July 11, 1932
(Taylor-Smith) ; 1881-1915A, 6 km. east, Amazoc, Puebla, July 20,
1932 (Taylor-Smith); 1916, La Colorada, Zacatecas (Smith and
Dunklc) , July 10, 1934; 1917-2009, 13 to 15 km. east Aguascalientes,
Aguas. July 23, 1934 (Smith-Dunkle) ; 2010-2014, Rancho Guade-
lupe, 42 km. west of Toluca, Mexico, August 4, 1932; 2015-2023,
Tepic, Nayarit (Taylor) ; 2024-2033, near Zapotiltic, Jalisco, June
24, 1935 (Smith); 2034-2058, near Chapala, Jalisco, July 2, 1935
(Smith); 2059-2066, near Uruapan, Michoacan, July 1, 1935
(Smith) ; 1878, near Tierra Colorada, Veracruz, July 16, 1932

Taylor: Frogs of Mexico 425

(Taylor-Smith); 1879-1880, 3 km. S. Hda. San Martin, near Zita-
cuaro, Mexico, August 4, 1932 (Taylor-Smith) ; 5600-5603, Tepeaca,
Puebla. August 12, 1936 (Taylor) ; 5604-5605, Hda. El Sabino,
south of Uruapan, Mich.oaean, August 5, 1936 (Smith); 5608-5617,
Uruapan, Michoacan, August 6, 1936 (Smith).

I have been able to examine a large part of the material of this
group present in the United States National Museum.

Diagnosis. The smallest Mexican representative of the cximia
group; usually some shade of green above and yellow below; dark
reticulation, enclosing cream spots on posterior and anterior sides of
thigh lacking; head proportionally small; a brown line from snout
passes through nostril to eyes, then continues diagonally low on the
side to groin, bordered above by a silver white line; lip bordered by
a dark line; area between the two dark lines on the side of head,
whitish, forming a stripe of irregular width to arm; front side of
tibia with some dim spotting, not or rarely bordered by a whitish
line on its dorsolateral surface. Trace of a web between fingers;
toes about one third to two-fifths webbed; the finger pads small but
distinct, one half to four fifths of the diameter of tympanum; a
tarsal fold; vomerine teeth between small choanae; eyelid narrower
than interorbital space; tibiotarsal articulation brouglit forward,
reaches to some point on eye. The horny nuptual pad present on
first finger a very light brown, but never a deep brown color.

Description of the species. The species is somewhat variable as
regards coloration and the character of the preserving fluid tends to
change somewhat the general appearance. Specimens preserved in
alcohol are usually some shade of gray or ultramarine; in formalin
they are often grayish-brown. Throughout its range the species
tends to break up into races, some of which will doubtless be named
eventually, when their ranges can be better determined. Four rather
distinct varieties are discernible. These vary somewhat in size; in
the size of the digital pads; the extent of the webbing on the toes;
the shape of the head, and in dorsal coloration. I can discern the
following varieties:

A. The typical form occurring in Puebla, Central Western Veracruz, Distrito
Federal, Me.xico, and po.ssibly also the borders of adjoining states. These
are characterized by the strongly defined lateral lines; usually a pair of
elongate spots in the sacral region, which may be extended forward or
broken into a second pair of spots, which are white-bordered, especially
on the medial edges.

B. A western striped form, best differentiated in Xayarit. In this the head
is usually less pointed and the dorsal stripes are wider, often reaching
forward to the eyelids; there is frequently a medial row of irregular dark

426 The University Science Bulletin

dots, and some small spots lateral to the dorsal stripes; often there is
little or no green coloring anywhere on body ; usually the thigh and tibia
have no, or only sparse, spotting abo\-e ; the range of this type includes
Jalisco, Na.yarit, Michoacan, and Morelos. In the latter region (Michoacan
and Morelos) many of the specimens are greenish and ha\'e the area be-
tween the dorsal stripe and the lateral stripe, \ery light (yellowish or
cream) and the medial and outer lateral rows of small spots are wanting.
(See plate XL\% figs. 1, 2, 4, 5. 9, 10.)

C. Specimens from Zacatecas and Aguascalientes have the hind limb \ery
heavily barred with brown dorsally. They are somewhat smaller, but
have much of the general appearance of the typical specimens. Sometimes,
howe\"er, the stripes reach the eyelids and certain ones .show spots lateral to
the dor.^^al stripes. (See plate XLVI, figs. 3, 6. 7. 8.)

D. The most striking variant is represented by two siiecimons from Durango
(USNM, No. 14083). These have a brownish ground color; one has large,
irregular, deep-biown s]30ts co\ering the dorsal surface of the body and
limbs; the other a pair of cur\ing shoulder spots with other scattered
spots and fine scattered flecks o\er the dorsal surfaces of the body. These
specimens are badly shrivelled; however, the pads on fingers are distinctly
larger than those on specimens from Xaj^arit.

These variants appear different to the eye. However, with the
study of the material avaihable to me I am unable to make a satis-
factoiy separation of these variant forms into groups that warrant
subspecific designation.

All seem to agree on having small choanae; relatively large groups
of vomerine teeth, and the same essential color pattern on sides of
head and body.

The figures given by Baird, United States and jNIexican Boundary
Survey (plate 38, figs. 8-10) are poor for detail, especially as regards
the markings on the side of the head, the general characters of the
roof of the mouth and the characters of the pads; and the general
habitus of the body (in fig. 8).

I am at a loss to account for the apparent absence of H. eximia
from Guerrero. Kellogg {loc. cit.) lists specimens collected by Hans
Gadow from Tacubaya, Guerrero. Since the postal directories of
1892 fail to list such a locality for ( iuerrero, I suspect that the state
is incorrect. This name occurs in Distrito Federal, where Gadow
collected, as well as in certain other states which I believe he did not
visit. (Tamaulipas, Campeche, Yucatan, Tabasco).

Hyla ewphorhiacea Giinther

Hyla euphorbiarea Gunther, Cat. Batr. Sal. British Mus., Feb. 12, 1859, p. 109. jil. X.
fig. C (type description; type locality, Cordova [Veracruz]; Cordilleras [of Mexico?] and
Mexico, four specimens and a skeleton).

Hyla eximia ? Peters, Monatsh. Akad. Wiss. Berlin, 18(;i1, p. 880 (part.). (It is possible
that all the specimens mentioned are true H. eximia. However, H. euphorbiacea is placed in

Taylor: Frogs of Mexico 427

synonymy.) Bowlenger, Cat. Batr. Sal. S. Ecuad. British Mus., 2(1 Ed. 1S82 (part., speci-
mens, types of H. euphorbiacea) ; Giinther, Biologia Centrali-Anuricana. Ropt. Batr; June,
1901, p. 2G1-262 (part.) Brocchi, Mis.sioii Scientifiqne au Mexique et dans I'AiiieriQue Centrale,
pt. 8. 2d section, livr. 1, 1881, pp. 32-33 (part.); Kellogg, Bull. U. S. Nat. Mus., No. 160,
1932, pp. 164-167 (part.) (all or part of the Oaxaca specimens — certainly rSXM. Xo. 47908).
f Hyliola bocourti Mocquard, Nouv. Aich. Mus., (4) T. 1, 1899, pp. 341-342 (type
description; type locality, "Haute Vera Paz," Guatemala; maxinunn length, 40 mm.; the
posterior face of the thigh is brown with yellowish white spots).

A series of nine small hylid frogs collected by Dr. Hobart M.
Smith near the city of Oaxaca make evident the necessity of reviv-
ing the name Hyla euphorbiacea Giinther. These specimens agree
with Giinther's brief description and figure. While obviously of the
eximia group, and related to Hyla eximia, they differ from this
species in the pigmentation and coloration of the posterior femoral
regions, and somewhat greater webbing of the toe, and in having a
broader head. Hyla regilla listed by Brocchi (loc. cit.). plate XIII,
figs. 2, 2a, cannot be H. bocourti ( euphorbiaecea) , since the dorsal
coloration and the posterior thigh coloration shown is different from
that given in Mocquard's description of Hyliola bocourti. Figure
3 of this same plate is labeled Hyla eximia var. euphorbiacea. This
is a specimen lacking black lines on the back and may or may not
be Hyla euphorbiacea, since certain eximia lack evidence of these
spots or lines, and they are frequently present in euphorbiacea.

Hyla bocourti Mocquard, Bull. Soc. Philom.. Paris, (9) T. 1, 15,
is very probably a true Hyla eximia of the Tepic form shown in this
paper (plate XLVI, figs. 1, 2, 4, 5).

Diagnosis. A member of the eximia group, related to Hyla eximia
Baird, but somewhat larger with a broader head and the posterior
part of the femur strongly marked with blackish pigment enclosing
round or irregular cream markings. Upper labial edge dark gray,
bordered by a cream line; greenish, or grayish-green above with
elongate spots which may be absent or obscured; vestige of web
between the fingers, with traces of dermal fringes on fingers; toes
with webs extending about a third of their length, with strong
dermal fringes; a well-defined tarsal fold; a large inner and smaller
outer metatarsal tubercle ; largest disks on fingers about two thirds
to three fourths the area of tlie tympanum; a fold on breast. Male
with vocal sac; tibiotarsal articulation reaches to about middle of
eye; length of eye slightly longer than its distance from the nostril,
one fourth shorter than length of snout; interorbital width about one
third greater than width of upper eyelid; vomerine teeth in two
slightly diagonal series between the choanae.

Description of the species. (From EHT-HAIS, No. 3134, neai"

428 The University Science Bulletin

Oaxaca, Oaxaca, Mexico, August 4, 1935. Hobart Smith, collector;
one of a series of 9 specimens.) Top of head nearly fiat; the depth at
tympanum, 4.5 mm., at nostril, 3.2; eyes moderate, projecting some-
what, tlieir length a little greater than distance to nostril, but much
shorter than snout; tympanum rather distinct, its diameter a little
greater than half the length of eye opening; upper eyelid about two
thirds as wide as interorbital distance; snout projecting moderately
beyond mouth. Canthus rostralis more or less distinct, rounded or
slightly angulate, the loreal region strongly oblique, not or slightly
concave ; vomerine teeth in two groups more or less closely approxi-
mated, lying between the large choanae, separated from them by a
distance slightly less than length of a single group; tongue nearly
circular, slightly notched behind, free for a little more than one
fourth its length; a large median vocal sac; skin above and on sides
smooth; abdomen and most of ventral surface of the femur with
large granules; chin with very minute granules; breast nearly
smooth, the granulation can scarcely be discerned; a part of the
posterior part of femoral region granulate; arm with a few granules
or tubercles on the ventral surface; fingers with well-developed disks,
that en first finger small, scarcely wider than chgit; that on third
finger more than two thirds size of tympanum ; first finger reaching
a httle beyond the subarticular tubercle of the second; latter reaches
the disk of fourth; the fourth reaches the disk of third; a vestigial
web between the fingers and a trace of a dermal fringe on sides of
digits; inner palmar tubercle large, prominent, rounded anteriorly;
median and outer partly confluent, forming a large tripartite pad;
subarticular tubercles, large, rounded, save the proximal tubercles on
two outer fingers, which are scarcely larger than supernumerary
tubercles on palm; hand, to tarso-metatarsal joint, shorter than the
tibia; when limbs are folded at right angles to body axis the heels
overlap about one millimeter. Inner metatarsal tubercle large, sa-
lient, its length in its distance from tip of first finger about 2.5 times;
toes about one-third webbed, the webbing exi:ending to or a little
beyond the second subarticular tubercle on the three outer toes, to
the single subarticular tubercle on the first and second; well-defined
dermal fringes on toes extending to the disks.

Color. Above green (silvery gray in alcohol) with two elongate,
irregular dark blotches beginning about middle of body and extend-
ing to groin; a dark narrow black bar extends from snout through
nostril to eye, silver-edged above; a broad, dark-blackish stripe,
silver-edged above, begins behind eye, including most of the tympa-

Taylor: Frogs of Mexico

429

num, then passes above arm; it then turns down, passing along the
sides of body, rising again posteriorly; the lower part of the stripe
is grayish posteriorly on its lower side and fades to a lighter shade
toward the ventral surface; a few small, indefinite, lighter spots on
sides; two or three larger yellowish spots in groin; upper lip bordered
with a narrow gray line continuous in front and bordered above with
a narrow cream or silvery line which is scarcely distinguishable
from the lighter area in the loreal region, which continues back a
little distance behind the angle of the jaws; outer edge of arm with
a light-edged, broad, dark stripe; an irregular darker blotch on hand;
palm lacking pigment; a dim darker bar crosses femur distally; one
or two similar bars on tibia; anterior surface of femur with a dim
dark stripe, light-edged above; a large blotch on foot near heel;
underside of foot and heel with some dark pigment; chin and other
ventral surfaces creamy white (vocal sac more or less darkened in
males) ; the posterior and anterior surface of femur blackish or
brownish with well-defined yellow-cream spots, round or irregular
in shape; the front side of the arm with none, or only very dim,
markings.

Measurements of Hyla euphorhiacea Gunther

Number.

Sex

Snout to vent

Head length to angle of jaw

Head width, greatest

Diameter of eye

Diameter of tympanum. . . .

Length of snout

Interorbital distance

Upper eyelid

Arm

Hand and longest finger. . . .

Leg

Tibia

Foot and tarsus

Largest finger pad

126

USNM
47908

3133

3134

&

cf

cf

9

35

37

32.2

33

10

11.5

10

10

12.2

12.9

10.7

10.5

2.5

4

3.5

3.6

2.1

2.2

2.1

1.8

4.9

5

4.4

4.4

3.65

3.8

3

2.9

2.8

2.4

2.5

2.4

20.2

21.5

19.8

20

9.9

11

8.6

9.1

55

60

51.5

50

16.6

19

15.2

14.2

25

27

23.2

22.3

1.9

2

1.5

1.3

Variation. The series of ten specimens vary in the presence of
visible dark spots on the sacral region. In some they are totally
obscured, in others they may be more distinct and there may be one
or more small spots farther anteriorly; the loreal region below the

430 The University Science Bulletin

black bar, and the region under eye and in front of tympanum may
be lighter or darker, but posteriorly the light color is prominent and
strongly delimited by darker color. The groups of vomerine teeth
vary in their proximity, sometimes being closely approximated, in
others rather widely separated; usually they reach both anterior
and posterior level of choanae.

Remarks. This lot of specimens was collected by Hobart M.
Smith, very near the city of Oaxaca, Oaxaca. In his diary he states:
''This hyla has a different call froiii H. cximia. The green color on
the back is about the shade of that in the leaves of a water lily.
The dark stripe on the side is brownish. The groin and concealed
surfaces of the femur and tibia are, in the lighter areas, yellowish
with a fine brownish reticulation. The vocal sac of the male is
light yellow with fine brown stippling. One copulating pair was
taken. They were extremely wary and were captured with difficulty.
All were captured in a rain pool in a cornfield."

This species differs from the typical H. eximia in having a some-
what larger size, and broader head; slightly more elongate fingers,
with the vestigial web more pronounced and with somewhat larger
disks; the webbing extends a slightly greater distance on the toes,
and the posterior coloration of the thigh is wholly different.

Giinther states: "M. Salle generally found it resting on the large
euphorbiaceous plants."

Hyla cdrdenasi sp. nov.

(Plate XL\II; fig. 2)

Holotype. USNM. No. 84403. Puebla. Puebla, Mexico, Septem-
ber, 1919. H. Ruano, collector. (Field No. 3, Comision Geografica
Exploradora de Mexico.)

Paratype. EHT-HMS. No. 3963, near Rio Frio, Mexico, July 31.
1932, E. H. Taylor, collector.

Diagnosis. A member of the eximia group. Eyelid wider than
the interorbital distance ; eye longer than snout in adult ; tympanum
more than half diameter of eye; head wider than long; diameter of
largest finger pads slightly greater than half the diameter of the
tympanum; pads subtruncate; web rudiment between first three
fingers, none between outer fingers; a thick, well-defined tibiotarsal
fold ; when limbs are folded, heels touch ; tibiotarsal articulation
reaches slightly in front of eye; tongue with fine raised papillae;
vomerine teeth in two small groups close together, lying directly
between choanae, but separated from them by a distance greater
than a single group.

Taylor: Frogs of Mexico 431

Description of type. Head very short, the snout as deep as its
length or approximately so, extending somewhat beyond mouth;
length of the snout distinctly shorter than length of eye; eyelids
distinctly wider than interorbital distance; diameter of tympanum
more than half the length of the eye; distance between nostrils
equals distance from eye to nostril; tympanum separated from eye
by a distance equal to three fourths its diameter; tongue sub-
circular, very slightly emarginate behind, covered with numerous
salient papillae; choanae moderately large, separated by a distance
greater than that between nostrils, but not greater than distance
between outer edges of nostrils; groups of vomerine teeth rather
narrowly separated, about size of choanae, not reaching anterior
level of choanae, but extending half their length behind hinder level ;
r-kin of dorsal surface quite smooth, on sides growing somewhat
pustulate or granulate toward ventral surface; chin and throat with
small indistinct granules; remainder of the ventral surface of the
body strongly granulate, the granules unequal in size; most of
proximal half of ventral surface of femur strongly granulate; distal
half smooth ; a few enlarged granules below anus, the median ones
bordering a distinct groove; some indistinct granules on the ventral
surface of arm ; a tliick fold behind eye covering upper edge of the
tympanum and extending back toward arm insertion, not folding
down behind tympanum; an area of thickened skin somewhat back
of the angle of the jaw.

Tips of fingers strongly dilated, the tips more or less truncate,
the essential portion of the pad (anterior to groove) practically
twice as wide as deep, the free edge of skin above the tip scarcely
evident ; a mere vestige of a web between first three fingers ; scarcely
or not indicated between the two outer fingers; the diameter of the
largest pad on fingers as great as that of tympanum, but area is
smaller than area of tympanum; distal subarticular tubercles large,
rounded, not divided; numerous supernumerary tubercles on palmar
surface. A large flat tubercle on ventral surface of the base of the
first finger, and a large flattened palmar tubercle, partially divided
anteriorly, on posterior part of palm ; narrow more or less con-
tinuous skin folds on outer edges of fingers, not or scarcely indicated
on inner edges; no folci on ventral surface of forearm; tibiotarsal
articulation reaching to about anterior edge of eye; a well-defined
tarsal fold becoming thickened posteriorly. Toes about one-third
webbed, the web continued on sides of toes as a narrow fringe, save
there is none on outer edge of outer toe; terminal disks smaller than
on fingers, the essential pad nearly twice as wide as long, the tips

432 The University Science Bulletin

rather truncate; all subarticiilar tubercles well developed, the distal
ones not noticeably larger than the proximal ones; supernumerary
tubercles on fingers low, rather indistinct; a strong oval, inner
metatarsal tubercle; a small indistinct outer tubercle anterior to
level of the anterior edge of inner.

Color in alcohol. Above a uniform purplish or lavender-gray
(probably a shade of green in life) with no evidence of darker
markings; no trace of a light line from eye along side; upper lip
scarcely lighter than remainder of side of head and no light mark
continued to arm from eye; lighter about insertion of arm; no light
mark above anus; posterior surface of femur heavily stippled or
powdered with brown; chin of female heavily pigmented; ventral
surfaces of hands lacking pigment; feet with some pigment; ventral
surface of body dirty-white, lacking pigment.

Measure77ients in mm. of the type and paratype of Hyla cdrdenasi.
USNM, No. 84403, and EHT-HMS, No. 3963, respectively: Snout
to vent, 39, 22; head length to jaw angle, 12, 8.2; head, greatest
width, 14, 8.2; diameter of eye, 5, 3; length of snout, medial, 4, 3.4;
depth of snout in front of eye, 4.1, 3.2; diameter of tympanum, 2.6,
1.6; interorbital width, 3.4, 2; upper eyelid, 3.4, 2; arm, 23.7, 12.2;
width largest finger pad, 2.5, 0.9; lengtli of leg, 67, 35; tibia, 21, 11;
foot and toes, 30, 15.2; largest toe pad, 1.5, 0.9.

Variations. The paratype is a young specimen. It will be noted
from the dimensions presented that the proportions vary between
young and old, the head being proportionally narrower and the
limbs proportionally different from those of the adult. The skin is
thickened, minutely corrugated. In life the color was a deep bluish-
green; in preservative it is ultramarine.

Remarks. The paratype was collected in the pine forest near
Rio Frio, Mexico, at an elevation of about 3,000 meters. It was
hopping about on the forest floor. The type is an adult female, the
ovaries filled with large eggs.

The relationship of the species appears to be closest to Hyla
lafrentzi, from which it differs in the details of markings ; in lacking
the pronounced dark and white lines on the side of the head and
body; in the somewhat wider and more truncate pads on the fingers;
in having a shorter^ thicker snout, and in the heavy pigmentation of
the throat of the female. Other differences are evident on a com-
parison of the descriptions and figures.

In size the species is intermediate between eximia and lafrentzi.

The species is named for General Lazaro Cardenas, president of
the Republic of Mexico.

Taylor: Frogs of INIexico 433

Ihjln lajrcntzi Alertcns and ^Yoltcl•st<)l■ff.

(Plate XL\'1II ; ties. 1,2)

Hylu lajrcntzi Meiti-ns ami \\ ..ItiTstcntf. Zo,il. Aiiz., H. 84, N i. D/ld. August i:,. Wti'.K jip.
23o-241 (t.\-pe tlespription ; type localitx', Desitrto de los Leones, in iiinuiitain forest, 3,000
meters elevation, near Mexico City, Distrito Fiderai. T.\pe in Magdehurs Miis. No. 49/27; K.
Lafrentz, cullector, December I84 1927); I-afrentz, Blatt. Aquar-Terrar. Kunde, 38, 1927, p.
322.

Hyla graciUpes Kellogg, Bull. U. S. Nat. Mus., No. IGO, 1932, pp. l,-.3-154, 1G8-170
(part.).

This species is represented in the collection by EHT-HMS. Nos.
3958A, 3959-3962, near Vigas, Veracruz, Mexico, July 13, 1932,
Taylor and Smith, collectors; and Nos. 5978-5991, Lake Zcmpoala,
near Tres Marias (Tres Cumbres), Morelos, 3,300 meters elevation.
(Lake Zempoala is about thirty kilometers from the type locality.)
I have examined two Michigan Museum specimens, Nos. 5304 and
48065, Guerrero, Hidalgo.

Diag7iosis. The largest species of the H. exiniia group, attaining
a known maximum size of 50 mm. head-body length; a vestigial
web between fingers; toes between one half and two thirds webbed;
fingers and toes with rather small terminal disks, those of fingers
the larger; those of outer fingers about half the area of the tym-
panum; large nuptual callosity at base of first finger, covered with
a deej) l)rown horny excrescence in males; tympanum one half to
two thirds the length of the eye; eye somewhat shorter than the
snout, the interorbital space somewhat wider than upper eyelid;
tibiotarsal articulation reaches tip of snout or somewhat beyond; a
distinct tarsal fold. Greenish above, the ventral surfaces yellowish,
save the chin may be dark in males; paired black spots edged with
white on sacral region and a dark blackish streak through eye to
side, edged above with white.

Description of species (from EHT-HMS, No. 1814, collected near
Vigas, Veracruz, July 13, 1932). Head rather thickened with a
depth at jaw angle of 6.5 mm., sloping forward to nostril, where the
depth is 3.8 mm., then sloping abruptly down to the mouth; the
snout very slightly rounded, in lateral profile, and projecting slightly
beyond mouth; canthus rostralis distinct to nostril, slightly rounded
rather than angulate; eye moderately prominent; the interorbital
width slightly less than width of an upper eyelid; length of eye less
than the length of snout^ nostril nearer the eye than to the median
anterior edge of mouth; tympanum rather large, its diameter a
little more tlian half the length of the eye; pupil of eye horizontal;
loreal region sloping obliquely to moutli, slightly concave, the lines
of the canthus nearly straight, when extended intersecting at tip of
snout or slightly beyond.
28—4141

434 The University Science Bulletin

Vomerine teeth in two rather small, somewhat conical groups,
lying wholly between the choanae, separated by a narrow distance,
and separated from the choanae by a distance slightly less than
width of a single group; choanae smaller than a single tooth group;
tongue subcirculai', large, emarginate posteriorly, free for a little
less than one third its length; vocal sac medial, single, evident on
chin by ample folding of the skin, its elongate openings lying lateral
to the posterior part of tongue.

A well-defined supratympanic fold, overhanging the tympanum
somewhat, runs back and down to a point somewhat back of the
insertion of the forearm; a glandular area behind the angle of the
jaws which is rounded on the posterior border; an indefinite glandu-
lar area on anterior proximal portion of upper arm. The skin above
is practically smooth (minutely corrugate) ; sides indistinctly granu-
lar or areolate, becoming more distinctly granular low on sides; en-
tire abdomen and underside of thigh (largely) with large distinct
granules, while those on breast and distended membranes of the
throat, smaller, with still smaller intercalated granules; a heavy
skin fold across the pectoral region, in which the granules are less
distinct; anal region granular; underside of arm granulate, but
remainder quite smooth; skin glandular in the supranal region.

A distinct vestige of a web is present between the fingers, and
evidence of a thickened dermal fringe on edges of digits extending
to the disks. Disks distinct, rounded, that on the third finger
largest, about one half the area of the tympanum, that on first finger
distinct, but scarcely wider than digit; a large nuptual callosity on
inner side of first finger extending to the disk, covered with a dark
horny excrescence (perhaps present only during breeding season I. A
very large palmar tubercle present on base of first finger, its inner
outline obscured by the nuptual callosity; medial and outer palmar
tubercles confluent posteriorly, separated by a groove anteriorly;
subarticular tubercles large; surface of palm with large, irregular
granules or tubercles; foot moderately elongate, the part from the
tarso-metatarsal articulation to tip of longest toe a little shorter
than tibia; a large inner metatarsal tubercle, its length contained
in its distance to tip of first toe one and two thirds times; outer
metatarsal tubercle small, rather indistinct. Toes somewhat more
than half webbed, the webs continued to tlie terminal disks as
nanow, thickened dermal fringes; the proximal subarticular tubercles
on the two outer toes small, others large; foot witli numerous in-
distinct subarticular tubercles; a clearly-defined tarsal fold; the

Taylor: Frogs of Mexico

435

dermal frino;e on the fourth toe continued a little behind proximal
subarticular tubercle.

Color in life. Upper parts dark green to olive-green, somewhat
lighter on the sides (bluish in alcohol). A dark bar from nostril to
eye and a dark grayish line bordering the lip, darkest on its upper
edge, and bordered above by a cream line, which terminates below
angle of jaw; a more or less dense black stripe, edged above with
cream, runs from eye along the supratympanic fold, and is continued
on the side of the body as a very narrow, scarcely discernible, dark
line to groin, delimiting the greenish color of back and sides.

A pair of dark, light-edged spots on the posterior third of the
body, with two or three spots on back between femora; a single in-
distinct dark spot on distal surface of femur; a narrow, liglit-edged,
dark line on outer (anterior) face of tibia, and a similar line on
outer (posterior) edge of the tarsus, continued along outer toe; the
dark line merges into the coloration below it ; glands above anus,
cream; posterior surface of femur and concealed ]iart of tibia with
a sparse, uniform peppering of black; a few indistinct darker flecks
on toes; ventral coloration cream to flesh, the vocal sac grayish-
purple with the granules cream; ventral surface of foot sparsely
pigmented; a dark line with a light border above on outer side of
arm and hand.

Measureinent.s (in mm.) of Hijla lafrcntzi Mertens and Wolter.storff

Number

Sex

Snout to vent

Snout to eye

Head length to angle of jaw . ,

Head width, greatest

Diameter of eye

Diameter of tympanum

Eye to nostril

Depth of head in front of eye

Heels overlap

Arm

Hand

Leg

Tibia

Foot and tarsus

5980

9
50

5.5
15
15.9

5

3

3.5

4.7

3.7
29.2
15
85

27.2
37.1

5984

5986

9
44

5.2
14.6
15.2

4.7

3

3.7

4.5

3
28
12.1
75.5
23
33.5

&

41.2
5.9

13.2

15
4

2.6
3.35
4.5
3.5

23

12.1

68

22

32

5978

43.2
5.8
13

14.6

4.3

2.85

3.6

4.5

2.7

23.6

12.1

68.3

21.5

31

5991

3960

d'

40
5.6

12

13
4.1
2.35
3.1
4.5
2.5

24.6

12.3

07 . 5

21

29

39
5. 5
12.3
14.1
4

2.3

3.3

4.4

3

24.5

11.6

64

20.2
28.9

436 The University Science Bulletin

Variation. Specimens from Lake Zempoala, Morelos, from a
higher elevation show a sliglit difference in the depth of the head,
and the tip of the snout is slightly more elongated.

Remarks. The specimens collected near Vigas, Veracruz, were
found about a small rain pool beside the highway during the morn-
ing. The males were calling. Those taken at Zempoala were calling
most of the day. A single pair was found clasping. A few im-
mature tadpoles, presumably of this species, were found in small
pools in the bog near the lake edge.

While this species resembles certain specimens of Hyla eximia in
general characteristics, its much greater length and bulk, the longer
legs, proportionally wider head, and the fact that the range of H.
lajrentzi is within that of H. eximia, should preclude the possibility
of their being regarded as the same species.

Hyla ivrightorum sp. nov.

(Plate Xl.Vn, fig. ].)

Hyla eximia Yarrow, Bull. U. S. Xat. Mus., No. 24, 1882 (part.) (specimens from New
Me.Kico) ;King, Copeia, 1932, No. 2, p. 99 (Mormon Lake, Arizona); Wright and Wright,
Handbook of Frogs and Toads, Ithaca, N. Y., 19.33, pp. 118-119, plate XLIII (an e.xcellent
series of photographs of live specimens with a description, and notes on voice, breeding, and
habits (Texas, New Mexico, Arizona) ; Stejneger and BarV)our, Check List North Amer. .Amph.
Rejit., 3d Ed., 1933, pp. 34, 35 (part.).

Hyla gracilipes Kellogg, Bull. U. S. Nat. Mus., No. 160, pp. 154, 168-170 (part.) (speci-
mens listed from Chihuahua) ; Stejneger and Barbour, Check List North Amer. Amjih. Kept..
3d Ed., 1933, p. 35 (part.).

Holotype. No. 79141, Museum of Zoology, University of Michi-
gan. Eleven miles south of Springerville, Apache county, Arizona,
U. S. A.

Paratypes. USNM, Nos. 26605-26609, Meadow Valley, Chihua-
hua, Mexico; USNM, No. 9338 (2 specimens), Santa Fe, New
Mexico, U. S. A.; MUMZ, Nos. 79141, 3 specimens; 11 miles south of
Springerville, Apache county, Arizona, August 13, 1935; 79143, 4
specimens; David Lee Lake, southwest of Luna, Catron county, New
Mexico, August 6, 1935, 8,000 feet; 79142, 3 specimens; 26 miles
north of Luna, Catron county. New Mexico, 8,100 feet, July 31.
1935; 75734, l^/o miles northwest of Miller's peak, Huachuca moun-
tains, Arizona, August 16, 1933.

Diagnosis. A member of the Hyla eximia group, but differs from
typical H. eximia in a larger size, longer legs, the heels overlapping
one or two millimeters when limbs are folded at right angles to the
body; the tibiotarsal articulation reaches the tip of snout or slightly
beyond; the head is less pointed, and proportionally wider; the first
finger is proportionally longer. The posterior half (or more) of the
edge of the lower jaw is darkly pigmented; the posterior side of fe-
mur lightly and evenly pigmented.

Taylor: Frogs of Mexico 437

Description of the type. A medium-sized member of the Hyla
eximia group. The snout is rather truncate or hknitly conical, with
the canthi more or less distinct but rounded; the line between eye
and nostril somewhat concave, sloping obliciuely from canthus to
edge of lip; diameter of eye somewhat greater than distance of eye
to nostril, and equal to distance of nostril to middle of upper labial
border; nostrils below edge of canthus, the distance between them
about equal to their distance from eye; the area about nostril
slightly elevated, and a slight, shallow groove present between
nostrils; diameter of the tympanum is contained in the diameter
of the eye slightly more than 1.5 times; the distance between the
tympanum and eye about .65 of diameter of tympanum.

Tongue broadly cordiform or subcircular with a very sliglit median
emargination posteriorly; free posteriorly for two fifths of its length.
Ill iiiak's The openings of the single vocal sac are lateral to the
tongue and much elongate; tongue papillae not prominent; the
raised prominences bearing the vomerine teeth are large, placed
slightly diagonally and closer to each other than to choanae; they
arise near anterior level of the choanae, but do not reach their
posterior level. The openings of the mucous glands form a con-
tinuous groo^'c anterior to choanae; latter proportionally large.

A vestige of a web between first three fingers, but practically
obsolete between outer fingers; disks on the fingers moderate, only
a little wider than the toes, the widest one on outer fingers equal
in widtli to a little more than half the diameter of tympanum; first
finger reaching to a point halfway between the distal subarticular
tubercle and the terminal disk of the second; the distal subarticular
tubercles large, that on outer finger very slightly bifid on right
side (probably abnormally) ; a slight dermal fringe on the lateral
edges of fingers; fourth finger longer than second.

Legs elongate, the limb laid forward, the tibiotarsal articulation
reaches to the tip of the snout or beyond slightly; when limbs are
folded at right angles to body the heels overlap about two milli-
meters; terminal disks on toes not wider than digits, distinctly
smaller than finger disks; a well-defined tarsal fold; a prominent,
salient, inner metatarsal tubercle, its length in the first finger length
about two and one-half times; outer metatarsal tubercle distinct,
flattened, lying behind the anterior level of the inner tubercle;
inner toes webbed at base, the depth of web from one fourth to one
third the length of the outer toes; the web between the three outer
toes incised to a point one third the distance between the two

438

The University Science Bulletin

proximal siibarticular tubercles of the fourth toe; supernumerary
tubercles on palm and foot more or less distinct. (In males the
large tubercle at the inner part of the base of first finger is covered
with a corneous callosity, usually veiy light brown in color. ) Anal
flap rather wide, not especially modified; no axillary web; skin on
body relatively smooth, under magnification one observes minute
corrugations, more evident above eyes; a strong skin-fold across the
breast; ventral surface granulate, the granules on the anterior part
of abdomen largest, less distinct on throat and chin; granulations
prominent on median ventral, and to some extent, posterior part of
thighs; a rather thick but relatively indistinct fold above tympanum.
Color in alcohol. Above, on limbs and body, grayish-lavender
(probably some shade of green in life) ; two rather large and several
smaller spots in sacral region ; a narrow brownish line begins near
tip of snout, passes back through the nostril to eye; beginning
behind eye it involves tympanum and runs back low on the sides,
breaking up into irregular spots as it rises diagonally on the posterior
part of the side; limbs barred with brownish, two bars each on
thigh and tibia, several on foot; the tibial spots are continued onto
front face of tibia; upper lip, and to a lesser extent lower lip,
bordered with a narrow band of brown, narrowly edged with cream
or white; ventral surface dirty-white, immaculate; a well-defined

Measurements (in mm.) ol Hyla wrightuntm sp. nov.

Number

USNM
9338a

MZUM t
19141
Type

USNM
93386

USNM
26607

USNM
26605

Sex

Snovit to vent

Head length to jaw angle.
Head width, greatest . . . .

Diameter of eye

Diameter of tympanum. .

Length of snout

Iiiterorliital width

E.\elid

Foreleg

Hind leg

Tibia

Foot and tarsus

9

42

12.4

13.1
4.5
2.6
5.2
4
3.5

21.2

68

22

30

9

42

13

13.2
4.35
2.85
5.2
4
3.8

23.2

70.5

23

31.5

9

40

12*

12
4.3
2.3
5

3.6
3.2

22.6

62

20

27.5

9

37
13.2
12

4

2.5

5.4

4

3.5
21
60
20
26.2

34.2

12.2

11
3.5
2.1
5

3.85
2.9

18

55

18

23

* Head somewhat dried and distorted.

t Museum of Zoology, University of Michigan

Taylor: Frogs of Mexico 439

spot on anterior side of upper arm, and none or only a \-ague scatter-
ing of pigment on the chin and throat ; underside of feet and hands
with some pigment and lighter flecks; posterior and anterior part of
femur and groin region with an eciual distribution of fine, brown
pigment.

Remarks. Specimens of this species have been present in the
United States National Museum since 1874. Yarrow (North Ameri-
can Reptilia and Batrachia, Bull. U. S. Nat. Mus. No. 24, 1882, p.
172) lists No. 8508, 2 spec. Nutrias, New Mexico, and No. 9338,
2 spec. Santa Fe, New Mexico, June, 1874, H. W. Henshaw, collector,
under the designation Hyla eximia Baird. In Cope's Batrachia of
North America, Bull. U. S. Nat. Mus., No. 34, these specimens are
not listed under Hyla eximia, nor, so far as I can discover, is there
reference made to them.

In the Stejneger and Barbour, Check List of North American
Amphibia and Reptiles, 3d Ed., 1933, pp. 34-35, Hyla eximia is listed
from Mexico, Texas, New Mexico and Arizona. Hyla gracilipes
is likewise listed, p. 35. I have seen no specimens of the eximia
group from Texas, but if a form of the group occurs there, it may
belong to this species {irriqhtorjim). I have examined a series of
specimens of this species in the collection at Cornell University,
due to the kindness of Dr. A. H. Wright. These specimens were
collected near McNary. in Arizona, by W. C. Chapel, .Jr. ]\Ir. F.
Willis King Hoc. cit.) has published a short note on the species
based on specimens collected by him at Mormon Lake, Arizona.

This species is related more closely to H. rcgilla and H. lafrentzi
than to the typical H. eximia. From the former it differs in having
a smooth rather than pustular skin, and in having a longer leg, the
tibiotarsal joint reaching the tip of the snout or beyond, instead of
to the region of the eye. The webbing of the toes is somewhat less
and the diameter of the tympanum is greater than half the diameter
of the eye ; the toes and fingers are wider with somewhat wider pads.

From H. lafrentzi it differs in having tlie webbing between the
toes somewhat less with narrower fingers and toes, larger choanae, a
slioiter, blunter snout, somewhat deeper in front of nostrils. The
front edge of the tibia is heavily spotted with brown, instead of hav-
ing it blackish with a cream-white or silver line whicli is continued
to foot.

It is a species apparently adapted to scmidcscrt conditions.

The species is dedicated to Anna Allen Wright and Albert Hazen
Wright in recognition of their work in American herpetology.

440 The University Science Bulletin

PLATE XLVI

Hyla cxintia Baiid

Fig. 1. Ilyhi cximla Baird. EHT-HMS, No. 2021, $ ; Tepic. Nayarit,

Mexico. X i-

Fig. 2. Same. EHT-HMS, No. 2023, 9 ; Tepic. Nayarit, Mexico. X 1-
Fig. 3. Same. EHT-HMS, No. 1931. £ ; Aguascalientes, Aguascalientes,

Mexico. X 1-

Fig. 4. Same. EHT-HMS, No. 2018, <? ; Tepic, Nayarit, Mexico. X 1-
Fig. 5. Same. EHT-HMS, No. 2015, S ; Tepic, Nayarit, Mexico. X 1-
Fig. 6. Same. EHT-HMS, No. 1925, £ ; Aguascalientes, Aguascalientes,

Mexico. X 1-

Fig. 7. Same. EHT-HMS. No. 1924, $ ; Aguascalientes, Aguascalientes,

Mexico. X 1-

Fig. 8. Same. EHT-HMS, No. 1947, <5 ; Aguascalientes, Aguascalientes,

Mexico. X 1-

Fig. 9. Same. EHT-HMS, No. 2061, i ; near Uruapan, Michoacan. X 1-
Fig. 10. Same. EHT-HMS, No. 2064, S ; near Uruapan, Michoacan. X 1-

Taylor: Frogs of Mexico

441

PLATE XLVl

/

442 The University Science Bulletin

PLATE XLVII

Mexican Hylidae

Fig. 1. Hijla ivrightorum ^p. nov. Holotype. MZUM, No. 79141. 11 miles
.south, 8i)ringerville. Apache county, Arizona. U. S. A.; actual length. 42 mm.

Fig. 2. Hyla cdrdcnasi sp. nov. Holotype, USNM, No. 84403, Puebla,
Puebla, Mexico; actual length, 39 mm. (Spotting on back and limbs due to
injuries.)

Fig. 3. Hyla yracilipes, Cope. Cotvpe, USNM, No. 15318 9 : actual length,
22 mm.

Fig. 4. Same. Cotype, USNM, No. 15319 9 ; actual length, 23 mm.

Fig. 5. Same. Cotype, USNM, No. 15320 9 ; actual length, 21.8 mm.

Taylor: Frohs of Mexico

443

PLATE XLVn

^ if '

y-

444 The University Science Bulletin

PLATE XLVIII

Hyla lajrentzi Mertens and Wolterstorff

Fig. 1. EHT-HMS, Xo. 5978, $ ; Lake Zempoala, Morelos, Mexico; actual
Jength, 43.2 mm.

Fig. 2. EHT-HMS, Xo. 5980, 9 ; Lake Zempoala, Morelos, Mexico; actual
length, 50 mm.

Taylor: Frogs of Mexico

445

PLATE XLVm

/

/

^~. s

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV J June 1, 1938 [No. 20

A Cuiiti'ibutiun to the Taxonomy of the Subfamily
Issinae in America North of Mexico (Fulgoridae,
Homoptera)

KATHLEEN C. DOERING,

Department of Entomology, University of Kansas

Part II

Abstract: This jinper comprises the second part of a monograph deahng
with the taxonomy of the subfamily Issinae (Fnlgoridae, Homoptera) in
America, north of Mexico. In Part I * only the genus Dictyssa was discussed.
In this i>art a complete key to the genera, which number twenty-one in all,
is given and the following twelve genera have been studied in detail : Eiithiscia,
Hystcroptcrum. Dictyonia. Diclyssoiiia, Dictyonissus, Neaethus, Aliaodema,
I'lixcs, Tylana, Traxus, Thionia and Picumma. Of these twelve genera the
following contain only one species, being either monotypic or at least having
only one sjiecies that occurs north of Mexico : Enlhiscia, Dictyonia. Dictyssonia,
Misodcma. I'lixcs. Tylana. and Traxus. In the discussions of the remaining
five genera a key to the species in each genus is given. The genus Thiunia
contains seven species, Picumna two species, Hysteropterum seven species and
one variety, Dictyouis.sus two species, and Neaethus thirteen. The total num-
ber of sj^ecies studied is fifty-three. The following species are described as
new: Thionia omani, Picumma chinai, Hysteropterum jascomaculosum, Dic-
tyonissus riigrophilosus, Neaethus perlucidus, Neaethus sinehamatus, Neaethus
similis, Neaethus jacintiensus, N^eacthus curi<a)ninis, N^eacthus u)riformus, and
Neaethus dircrsus. Issomorphus niaculatus Melichar has been jilaced in the
genus Picumna.

Some of the older species have been redescribed and comiiarati\e notes
and diawings are given for all species except Hysteropterum morum Van Duz.
The male genitalia are described and figured for all species except the follow-
ing: I'lixes .■icutatus (not yet recorded from the United States), Thionia
omani and Thionia quinquata.

The characters of most \'alue for cla.ssification are the male genitalia, the
shape and position on the body of the tegmina, the size of the hind wings,
and the wing venation, including the fineness or abundance of reticulation.

* The I'liivcrsity of Kansas Science Bulletin. Vol. XXIW No. 17, I'.KiC,.

(447)

448 The University Science Bulletin

Key to the Gexera

1. Tiginiiia entirely covering abdomen or greater jiortion of it, iiarclunentlike or

vitreous, or a combination of both 2

Tegmina short and usually extending only partially over abdomen, except in
macropterous forms of some species which are always more or less hyaline,
parchment like, thick or opaque (Tribe Caliscelini) 16

2. (1) Hind wings either absent, rudimentary or else very narrowed with vein Sci

present and vein Cui branched or hind wings long, notched at apex, vein
Sci not jiresent and vein Cui not branched ; posterior tibiae with 2 to 5
spines or, entirely vmarnied 3

Hind wings present, narrow, not notched at ajiex. anal area small or else
rudimentary; jiosterior tibiae with 2 to 4 lateral spines; vein Sci absent

(Tribe Issini) 4

3. (2) Hind wings present, entire, with strongly marked notches at the joints of the

folds, anal area large; vein Sci absent; usually large insects varying from
5.5 to 8.1 mm (Tribe Thioniini) 6

Hin<l wings usually absent or rudimentary (a few species of Neaethus ex-
cepted); \ein Sci present (Dicti/ssoiua excepted); small insects, usually
under 5.5 mm (Tribe Hysteropterini) 7

4. (2) Frons deeply concave or perpendicular; cl.vpeus strongly deffexed and hori-

zontal Trarus Metcalf, p. 449

Frons not concave; cl.vpeus not so deeply inflexed or horizontal 5

5. (4) Tegmina rugulose with veins slightly elevated, finely reticulated; hind wings

small and narrow I'lixes Stal, p. 451

Tegmina not rugulose, with strongly elevated veins, larger reticulations; hind
wings half the length of tegmina, not visible when latter is in repose.

Tijlana Stal. p. 454

0. (3) Cubital vein of tegmen simple and costal region vertical to the body;

posterior tibiae with 2 spines TIdonia Stal, p. 450

Cubital vein of tegmen branched; posterior til>iae with usually 4 spines.

Picumna Stal. p. 471

7. (3) Tegmina more or less opaque or with \itreous sjiots and oblique bands 8

Tegmina vitreous entirely except Misodema and some species of Neaethus
parchmentlike 12

8. (7) Vertex not conically produced; posterior tibiae with one or more spines 9

Vertex conically produced; posterior tibiae without spines; tegmina not ex-
tending Ix'yond tip of clavus Eut)iiscia Van Duzee, p. 479

9. (8) Tegmina with vitreous patches or spots (Dictygsa fiisca excepted) not neces-

sarily narrowed at apices; posterior tibiae with 2 to 4 spines 10

Tegmina uniformly oiiaque with no vitreous spots or bands, narrowed and
rounded at apices; more or less thickly Viranched veins; posterior tibiae
with 1 or 2 spines Hyttteropteriim Amyot and Serville, p. 481

10. (9) Tegmina either as broad as long or approximately semicircular in shape,

I^artly opaque, usually with an obliqvie hyalin^ band across clavus and
cerium or some sort of hyaline markings; costal margin decidedly round-
ing 11

Tegmina oblong, costal margin nearly straight or if rounding with a distinct
bulla jjresent at base of each wing 14

11. (10) Tegmina semicircular in shape, not closely adpressed to body, veins of

corium forming irregular cells, some of which usualh' (not always) are

vitreous or light colored Dicti/ssa Melichar *

Tegmina practically as broad as long, held almost vertically, cells of corium

exceptionally few and large and distinctly angular Dictyonia Uhler, p. 496

* Key to species and descriptions of this genus in Part I of this paper in l'ni\-. of Kan.
Sci. Bull, Vol. XXIV, p. 424, 1936.

Doering: The Subfamily Issinae 449

12. (7) Upper surface hispid; vertex obtusely produced; posterior tibiae with 3

spines; tegmina with a fine network of coarse veins and main veins not
evident Dictyonii:s:us Uhler, p. 498

Upper surface not hispid ; vertex not produced or, if so, triangularly pro-
duced ; posterior tibiae with 2 or 4 spines 13

13. (12) Tegmina placed more or less vertically, close to body; posterior tibiae with

2 spines; head and eyes a.s broad as or broader than pronotum

Neaethus Stal, p. 501

Tegmina opaque, strongly inflated, gradually narrowed behind and with very
short clavus; head narrower than pronotum; posterior tibiae with four
strong spines Misodema Melichar, p. 534

14. (10) Tegmina broad, costal margin rounding 15

Tegmina decidedly oblong, approximately same length throughout, with costal
margin nearly straight Dictydea Uhler

15. (14) Tegmina with distinct bullae at outer angles of the cori\nn ; reticulation

finer Dictyssonia Ball, p. 536

Tegmina broadest just at middle; reticulation coarser Dictyobia Uhler *

16. (1) Body robust; narrow, lance-shaped tegmina, extending spearlike to or be-

yond apex of abdomen Danepteryx Uhler *

Body more elongate, tegmina not strap-shaped, usually covering only one
half of abdomen 17

17. (16) Head produced snout-like 18

Head not produced snout-like 19

18. (17) Fore and middle tibiae expanded Fitchiella (Fitch) *

Fore and middle tibiae not expanded Bruchomorpha Newman

19. (17) Tegmina oval and abbreviated; margin of vertex and front produced into

acute hornlike angles above eyes Osbornia Ball *

Tegmina more parallel, longer; no such acute angles above eyes 20

20. (19) Head large, including eyes, a little wider than base of closed tegmina ; front

rounded, prominent ; vertex short Aphelonema Stal *

Head, including eyes, narrower than prfjnotum or base of closed tegmina;

front triangular; vertex long and narrow Papagona Ball *

* Last seven genera will appear in part 3.

The Genus Traxus Metcalf, 1923

Metcalf, Z. P. The Fulgoridae of Eastern North America. Jr. of Eli.sha Mitchell
Society XXXVIII, 1923.

Comparative notes. This genus is separated from all other North
American genera by the deeply concave frons which is perpendicular
and the strongly inflexed horizontal clypeus, the basal margin of
which scarcely protrudes into the frons, except right at middle for
a short distance, where it appears rather bulbous as viewed from the
side. Other distinguishing characters are: the triangularly incised
vertex; rugulose appearance; tegmina thick with a roughened
texture, transversed by elevated longitudinal veins and a network
of less distinct veins at apex, costal margin not inflexed; clavus
very short, its apex only reaching halfway of the length of the
tegmen; hind wings small or rudimentary; number of spines on
hind tibiae two.

The wing venation seems to follow that of the genus Picnmna.
The general features are: vein Scj is lacking; veins Sc. and R
29—4141

450 The University Science Bulletin

united at base for a short distance only; apparently both veins M
and CUi are two-branched, although this is not always discernible,
since all the longitudinal veins, posterior to the median depression,
are lost in the uniform apical reticulation; vein Cu., follows the
claval suture; veins 1st A and 2d A united at apex to form the
characteristic Y-vein, although each are quite sinuate, pulled out
of a straight course by the concavity of the tegmen.

Traxus fulvus Metcalf

(Plates XLIX-LII-LIII-LV-LVI)

Metcalf, Z. P. The Fulgoridae of Eastern North America. Jr. of Elisha Mitchell
Society XXXVIII, 1923.

Comparative notci^. General color, grayish-tan with dark-brown
flecks and spots. The species is easily recognized structurally by
the subglobose, convex elytra with several prominent elevations
located as follows: the acute angle formed by the lateral margins
of the vertex and the frons where they meet above the eyes; on
each lateral half of pronotum a carinalike ridge following the eye
to the ventral side; on the mesonotum each lateral carina elevated
on a prominent boss; and a transverse hump, beginning on the
anterior half of the clavus and continuing across the corium to the
costal margin, back of which is a transverse concavity extending
across entire tegmen.

This species had been brought to the author's attention recently
by one or two colleagues, who were of the opinion that it was syn-
onymous with mixes scutatus Walker, a Mexican species, figured
and discussed by Fowler in the Biologia Centrali Americana. The
writer sent a specimen to Mr. W. E. China of the British Museum
for comparison with the type. In a letter to the writer, Mr. China
states: "This is nothing like Walker's species and does not belong
to the genus Ulixes. Indeed, it belongs to quite a different subfamily,
the Hysteropterinae. It appears likely to be a new genus. The
structure of the frons and clypeus is very unusual."

Male genitalia. Resembling the genitalia of Picumna in general
outline. The tenth abdominal segment (anal flap) seems distinctive
because of its ovate outline. It is a flattened tube, narrowed con-
siderably at base, then rounding out through middle and from
thereon with its ventral margin extending caudad as a flat, round-
ingly pointed flap. The eleventh segment scarcely shows beyond
the dorsal margin of the tubular part of the tenth segment. It
bears a much shorter and thicker ventral stylus than is found in
other genera.

Doering: The Subfamily Issinae 451

Harpagones (genital styli of authors) visible externally as two
broad triangular plates which are about twice as long as width at
base. Each harpago as viewed from a lateral, flattened view is a
subquadrangular plate with its dorsal margin at about middle pro-
jected dorsad as an irregular flat extension terminating in a slender,
cephalad, curving hook and also bearing on its cephalic margin a
flat, recurved hook which bends caudad.

The aedeagus and theca are difficult to distinguish in this genus
as in Picumna or Tylana. Together they appear as a bilaterally
symmetrical, broad, tubular structure. From a lateral view the
theca shows its apical margin, broadly notched through middle, due
to its ventral region being greatly extended caudad as a slightlj
pointed lobe which bears on each side a rounded lip-like extension
lying in situ just posterior to each aedeagal hook, and its dorsal
region extended caudad into a bulbous knob. The apex of the
aedeagus apparently does not show beyond the theca. However,
two pairs of aedeagal hooks are conspicuous. One hook emerges
from the thecal notch, extends ventrad a short distance, then
abruptly bends cephalad and extends forward almost to the base
of the aedeagus. The second hook on each side also emerges at the
notch of the theca and is directed cephalad to a little beyond middle
of aedeagus. This hook is heavily sclerotized and is forked at its
middle into two sharply pointed, curved hooks, the dorsal one of
which is slightly longer than the ventral one.

Notes on distribution. Metcalf lists the type specimens as all
coming from Brownsville, Texas.

In the National Museum collection there is a pair which were
taken at Brownsville, Texas, in June by Paul Oman. In the Snow
Entomological collection at the University of Kansas is another
pair taken by R. H. Beamer at Cameron county, Texas, in August.
Apparently this is one of the rare species in this subfamily.

Location of types. Probably in the collection of Z. P. Metcalf,
Raleigh, S. C.

The Genus Ulixes Stal, 1858

(Plates XLIX, LII, LIV)

stal, Carolus. Bidrag till Rio Janeiro — Traktens, Hem. -Fauna, II, p. 67, 1858.
Fowler, W. W. Fulgoridae. Biologia Centrali Americana. Homoptera I, pp. 113-119,
1904.

Walker, F. Insects Saundersiana, Homop. p. 44, 1858 (Issus).

Comparative notes. Fowler characterizes this genus as follows:
Wings small or rudimentary, not notched at apex; frons perpendic-
ular, clypeus more or less inflexed; peculiar rough facies; strongly

452 The University Science Bulletin

raised venation; general shape convex or subglobular and tegmina
as a rule strongly convex (scutatus excepted) ; spines on hind tibia
two to four as given by Fowler in the Biologia Centrali Americana.
There are four in scutatus. Fowler in his original description for
intermedius states that there are four. The writer believes that
four is the correct number for the genus.

One of the most distinctive characters which the present author
notes is the short clavus, whose apex extends only slightly over one
half the length of the tegmen. Another is that the costal border is
not inflexed as in many other genera.

The general features of the wing venation as is found in Ulixes
scutatus are: a separate Sc^ vein lacking; veins Sco and R united
for a short distance at base, curving outwardly at this point so that
these veins as they extend parallel with each other to the apex, are
much nearer to the costal border than in most species; vein M fork-
ing into two branches about midway of the tegmen; vein Cu^ ap-
parently single-branched, its tip lost in the apical reticulation; vein
Cu2 following the anal suture; veins 1st A and 2d A united at tip to
form the characteristic Y-vein.

Key to Specie.s

la. Head shorter; tegmina more rounding at tip; frons perpendicular; clypeus strongly
inflexed Vlixes scutatus Walker.

lb. Head longer ; tegmina obtuse at apex ; f rons and clypeus gradually rounded under
head Ulixes intermedius Fowler.

Note. Only scutatus has been collected in this country to date.
Ulixes intermedius, however, was collected by Dr. E. D. Ball in the
Sierra Madre mountains, Mexico. His material was in the type
series of Fowler's and was sent back to him by Fowler. Doctor
Ball very kindly pointed out the differences of intermedius, which he
has in his collection, with a solitary specimen of scutatus, which the
writer sent to him for comparison. The differences which Doctor
Ball noted are incorporated in the above key, which the present
writer considered might be useful, insomuch as Doctor Ball has
mentioned that any Mexican material collected by him in these
mountains is almost certain at some time to turn up in the south-
west of our country.

The single female specimen of scutatus Walker, mentioned above,
has been compared by the writer with a single male specimen sent
to the writer for study from the British Museum collection. The
latter specimen was compared by Mr. China with the type. The
female specimen from the United States was identical to the male
specimen in body form and coloring in every particular with the

Doering: The Subfamily Issinae 453

following exceptions: the United States female specimen had the
clypeus more distinctly retreating than did the other one (see draw-
ing), and the female specimen, contrary to the general rule, was
smaller by one millimeter than the male. Since the male was col-
lected at Oajaca, Mexico, and the female at Yarnell, Ariz., this
difference might be accounted for by difference in range and the
structural difference by that of difference in sex.

Ulixes scutatus Walker

(Plates XLIX, LII, LIV)

Comparative notes. A large, broad species, the same or larger
than Tylana ustulata, measuring 7 to 8 mm. long and 5 to 6 mm.
wide across greatest width of tegmina. Greyish or greyish-brown
in color. Vertex and thorax uniformly brown, flecked with yellow
spots. Tegmina yellowish-tan, rather solidly flecked with brown.
Underside of abdomen yellow, sparsely marked with irregular brown
spots.

This species is readily distinguished from any other North Ameri-
can form by its broad, flaring tegmina, its short clavus, its inflexed
clypeus and the lateral carinae on pronotum being much nearer to
the posterior border of the sclerite than to the eye. In the inflexed
clypeus and also in the short clavus it perhaps more nearly re-
sembles Traxus fulvus than any other species. It is easily sepa-
rated from this species, however, by lacking the mesonotal bosses,
by having four spines on the hind tibiae instead of two and by the
general shapes of the bodies.

Male genitalia. No specimen w^as available to the writer for dis-
section. Only one specimen was obtainable for study. This was a
specimen from the British Museum, compared with the type by
Mr. China and loaned to the writer for temporary use. A drawing
of the external genitalia of this specimen is figured. Fortunately
the harpagones are spread apart so that the aedeagal structure can
be partially seen.

From this position it can be seen that the theca on the ventral
side extends to almost the apex of the aedeagus and on the dorsal
side extends slightly beyond, ending in a rather swollen knob. Aris-
ing from each lateral slit of the theca are two well-sclerotized
aedeagal hooks, the more dorsal one extending laterad on the outside
of the harpago and bending back cephalad so that its apex extends
under the rim of the last complete abdominal segment, the ventral
hook a long, bladelike structure, bearing a short lateral hook near

454 The University Science Bi'lletin

base and tapering to a finely pointed apex, which does not quite
reach the rim of the abdominal segment.

Notes on distribution. Fowler lists this species from Mexico and
from San Geronimo, Guatemala. In the National Museum at Wash-
ington is one specimen collected by Mr. Paul Oman from Yarnell
Heights, Ariz., in 1933. This one female specimen was compared
with a male specimen from the British Museum collected at Oajaca,
Mexico, which Mr. China compared with the Walker type and
loaned to the writer for study.

Location of type. British Museum, England.

The Genus Tylana Stal, 1862

Stal, Carolus. Rio Jan. Hemip. II, p. 67, 1862.

Comparative notes. Some of the outstanding characteristics of
the genus are: head, including eyes, somewhat smaller than thorax,
frons with three longitudinal carinae, not united at base, clypeus
inflated; pronotum with a raised anterior margin which extends
ventrad as a distinct carina which finally joins the posterior mar-
gin; scutellum tricarinate; tegmina obliquely truncate at apex,
with a very long clavus which extends approximately three fourths
the length of the tegmen, longitudinal veins very prominent and
rugulose, cross veins prominent at apex of tegmen, making a prom-
inent network; hind wings short, extending only halfway, not
notched at apical edge, the anal lobes not very broad; femora of
hind legs with two spines.

Tylana ustulata Uhler, 1876

(Plates XLIX, LII, LIII, LIV, LVI)

Uhler, P. R. List of the Hemiptera of the Region West of the Mississippi River, Including
Those Collected During the Hayden Explorations of 1873. Bui. U. S. Geol. Geog. Surv.
I, p. 354, 1876.

Comparative notes. There is only one species in the genus in
North America, north of Mexico. Therefore, the characteristics
pointed out as distinctive for the genus also are specially valuable
in separating this species from other members of the subfamily.
General color pitch-brown to pitch-black, with lighter markings,
the latter in most specimens making a light saddle across middle of
tegmina. Tylana ustulata is one of the largest species in the entire
subfamily, measuring 6 to 7 mm. in length from tip of head to apex
of tegmen. Superficially it closely resembles, in color and size,
Picumna maculata Melichar and Picumna chinai n. sp. From these

Doering: The Subfamily Issinae 455

it- can be distinguished by having the tegulae of the wings hidden
under the pronotum and the base of the wing somewhat twisted
so that the longitudinal veins appear to start immediately behind
the pronotal margin; by having prominent cross veins at apex of
clavus making a network; by having the lateral margins of the
vertex elevated and meeting the lateral carinae of the frons in an
acutely produced angle above the eyes; and by having the costal
margin broadly expanded and bent under for half the length of the
tegmen, at which point the costal margin is distinctly angulate, and
by having short hind wings, not over half the length of the tegmina.

Wing venation. Due to the bent-under costal margin of the
tegmen the subcostal vein does not show from a dorsal view of the
insect, and although this costal area is rather broadly expanded no
Scj vein apparently is present. Another peculiarity is that the
basal cell from which the longitudinal veins arise in other genera
seems to be obliterated or pushed so far forward that the veins ap-
pear to start just back of the pronotal margin, and lastly an ambient
vein is present, starting at apex of clavus and continuing around the
apical margin to the costal margin, where it ends at the angle half-
way down the length of the tegmen. All the veins run rather
parallel. Sc^ and R are united for only a very short distance, then
each extends as a single vein to the apical region, where they are lost
in the reticulation; M divides into two branches, each of which is
also lost in the apical reticulation; Cu^ is single branched; Cu., runs
along the claval suture ; 1st A and 2d A unite to form a prominent
Y-vein in the clavus.

Male genitalia. The tenth abdominal segment forms the so-called
anal segment or flap, whose apex lies closely against the harpagones.
Basally it is a broad tube which has its ventral margin extended
caudad as a broad flap. The lateral margin of the flap constricts
at middle, then rounds out slightly and finally tapers to the truncate
apex, which is slightly emarginate through middle. The eleventh
segment shows beyond the dorsal margin of the tubular part of tlie
tenth segment as a short ringlike segment, bearing a broad, pointed
stylus.

Externally, the harpagones (genital styli of authors) appear on
the ventral side of th^e body as two adjoining triangular plates,
whose combined width at base is about equal to their length. Each
harpago as viewed from a flattened lateral position (see drawing No.
4, plate LVI) is roughly quadrangular with its dorsocephalic angle
obliquely cut off and its dorsocaudal angle greatly prolonged dorsad

456 The University Science Bulletin

into a broad, sharply pointed projection, at base of which is a short,
broad, flaplike hook.

The aedeagus is bilaterally symmetrical, almost completely hidden
by the theca. AVith the theca the aedeagus from a ventral view ap-
pears to be a broad, capsulelike tube bearing two long tapering
sclerotized processes, whose apices extend beyond the base of the
genital tube. These processes are aedeagal hooks which extend out
from a notch in the theca at a point about one third of the way
from the apex of the aedeagus, then broadly curve ventrad a short
distance, after which they extend directly cephalad, to a point be-
yond base of the latter. The theca on the ventral side is truncate
posteriorly. On the dorsal side it is somewhat more membranous,
appears sunken lengthwise through the middle and has a spatulate,
lobelike extension just posterior to the base of the aedeagal hook
and is also expanded lobelike at apex.

Location of type ynaterial. United States National Museum,
Washington, D. C.

Notes on distribution. Uhler in his original description states
that this species inhabits Colorado and Arizona. Specimens are at
hand for study from the following places in Arizona: Sabino canyon,
Vail, Baboquivari mountains, Pima county, Miami and Tucson.
Specimens from Miami, Ariz., were taken by Dr. R. H. Beamer on a
tree-like form of Equisctum. One specimen was taken by R. H.
Beamer from Rodeo, N. M., in August, 1935.

The Genus Thionia Stal, 1859

Stal, Carolus. Novae quaedam Fulgorinem formae species que insigniores. Berl. Ent.
Zeit. Ill, p. 313, 1859.

Fowler, W. W. Fulgoridae. Biologia Centrali Americana, Homoptera I, pp. 113-119,
1904.

Melichar, Leopold. Monograph der Issiden (Homoptera) abh. K.K. Zool. — Bot. Ges.
Wien III, 1906.

Metcalf, Z. P. The Fulgoridae of Eastern North America. Jn. of the Elisha Mitchell
Society XXXVIII, 1923.

Comparative notes. The genus includes species of large or medium
size, varying from 5.5 mm. to 8.1 mm. The vertex is chevronlike
in shape, depressed through middle and with sharply elevated mar-
gins. The frons is elongate with elevated margins and three carinae
usually present [simplex excepted), the two lateral ones of which
have their posterior ends curving to meet the median one. The
costal area of the tegmen is not deflexed as in Tylana, nor horizontal
as in Picumna and Ulixes, but is in a vertical position, while the
rest of the tegmen is more or less slopingly horizontal to the body.

Doering: The Subfamily Issinae 457

The tegulae are inconspicuous, showing only at the extreme sides.
The longitudinal veins are prominent; cross veins are lacking for
the most part, but sometimes show as a faint network at apex of
the tegmen. Hind wings present and as long as the tegmina.
Posterior tibiae with only two spines.

Vein Sc in this genus is not branched, and, therefore, no costal
vein (Sci) is present. Vein Sc. and R arise from a main vein
trunk near the base of the wing as a single vein, but very shortly
separate into two separate veins which run parallel to apex of the
tegmen. Vein M divides near middle of the wing into two branches
and branch M^ ^ o "^'"^y or may not divide at apex. Cu, is simple,
Cu^ follows the anal suture. Veins 1st A and 2d A are united near
the apex of clavus to form the characteristic Y-vein.

The general color for the genus varies from tan to brown, some-
times unicolorous, but usually variegated with darker patches or
with small, dark-brown, round speckles or with both.

This genus resembles the genus Picumna Stal very closely, but is
easily distinguished by the bispinose hind tibia and vein Cu^ being
simple, while in the other genus the tibia is ciuadrispinose and vein
CUj branched.

HISTORY OF THE GENUS

Stal described the genus in 1859 in Novae Quaedam Fulgorinem
Formae Speciesque Insigniores, Berl. Ent. Zeit. III. Van Duzee gives
the logotype of the genus as longipermis (Spin.). In 1830 two
species, namely simplex and elliptica, were described by Germar, who
placed them in the genus Issus. In the same year Say described
bullata as a Plata. In 1905 Fowler described a species in Mexico
which he called naso. Recently a long series of this species has been
taken in our southwestern states. In 1908 Van Duzee added the
long-headed producta to the group and in 1923 Metcalf added
quinquata. In this paper one new species is being added, T. omani.

This makes a total of seven species, which represent in collections
the North American species of Thionia.

To this list may be added the two species of Melichar's (1906)
transversalis and ocellata, which the writer believes are not found in
this country, since the type locality was given merely as N. A. No
specimens are present in the National Museum collection and no
specimens have turned up in the numerous collections thus far
studied.

458 The University Science Bulletin

Key to Species

1. Vertex considerably broader than long 2

Vertex longer than broad 4

2. (1) Anterior margin of vertex rounding, not produced anteriorly beyond eyes;

tegmina narrow, not bullate, unicolorous simplex (Germar), p. 458

Anterior margin of vertex slightly angulate; tegmina bullate and broad at
base 3

3. (2) Vertex concave, lateral margins greatly elevated; tegmina not narrowed

considerably beyond middle elliptica (Germar), p. 459

Vertex not concave, lateral margins not elevated; tegmina greatly narrowed

and reflexed at apex omani n. sp., p. 461

4. (1) Head triangularly produced a considerable distance beyond eyes 5

Head produced only very slightly beyond eyes or not at all 6

5. (4) Head produced beyond eyes a distance more than or equal to the length of

the eye; a larger species, measuring 6.8 mm. to 8.4 mm. .. .naso Fowler, p. 463

Head produced beyond eyes a distance less than the length of the eye; a

smaller species, measuring 5.5 mm. to 6.8 mm producta Van Duzee, p. 465

6. (4) All six margins of the chevronlike vertex equal in length so that its latero-

anterior angle is posterior to the anterior margin of the eye; larger in-
sects, measuring 8 mm. for females qiiinquata Metcalf, p. 467

Lateral margin of vertex longer than the four posterior and anterior margins,
thus bringing the lateroanterior angle even with or anterior to the anterior
margin of the eye. Size, 6.4 mm. 1o 7.6 mm buUata Say, p. 468

Thionia simplex (Germar), 1830

(Plates L. LI, LIT, LIII, LV, LVI)

Germar. Thon's Ent. Arch., II, p. 51. Issus.

Dozier, Herbert L. The Fulgoridae or Plant -Hoppers of Mississippi, Including Those
of Possible Occurrence. Miss. Agri. Exp. Sta. A. & M. College, p. 105, 1928.

Comparative notes. This is the smallest species in the genus,
measuring 6 mm. to 6.5 mm. in length. It is distinguished easily by
color, being uniformly brownish-yellow without markings. Struc-
turally it is readily distinguished by the squarish vertex which is
broader than long; by the frons which is only slightly longer than
wide, narrowed between the eyes and bearing only one median
carina or sometimes with two lateral carinae just faintly indicated
on either side.

Male genitalia. The tenth abdominal segment forms the so-called
anal tube or flap. Basally it is a broad, fiat tube that has its ventral
margin greatly extended caudad into an elongate spatulate lobe or
flap which is truncate at apex. The eleventh segment shows beyond
the dorsal margin of the tubular part of the tenth segment as a short,
ringlike segment bearing a very short, slender stylus.

Externally the harpagones (genital styli of authors) appear on
the ventral side of the body as two broad triangular plates which
closely adjoin along their mesal margins. As viewed from a flat-
tened, lateral view each harpago is a broad, pear-shaped plate, nar-
rowed at base, greatly broadened across apex, where the dorsal.

I

Doering: The Subfamily Issinae 459

apical angle is extended dorsad in a slightly pointed, recurved proc-
ess, at the base of which is a broad, flat posteriorly projecting flap.
(See drawing 5, plate LVI.)

The aedeagal structure is similar to that of the genus Picumna,
being a bilaterally symmetrical, broad, tubular organ in which the
limits of the theca and aedeagus are hard to distinguish, since the
theca is sclerotized considerably and almost completely covers the
aedeagus. From a lateral view the theca can be seen to extend al-
most to apex of aedeagus. Its posterior margin is truncate on the
ventral side, but has its dorsal angle extended caudad as two proc-
esses, the dorsal one of the two being an elongate, slightly recurved
lobe, the ventral one a sharply pointed hook which bends somewhat
ventrad. The apex of the aedeagus shows as a thick lobe between
the processes of the theca and bears two long, narrow, pointed proc-
esses, which protrude, one on each side, from the lateral notches in
the theca and bend directly cephalad, extending as far forward as
the base of the aedeagus.

Location of type. Possibly in the Museum at Budapest.

Notes on distribution. This is a southern and eastern species.
The type locality is given as Kentucky.

A large series of this species was available for study from the
following localities: Alabama, Arkansas, Florida, Mississippi, North
Carolina, South Carolina, and Virginia.

Dozier (1928) adds the following states in which it has been col-
lected: New Jersey, Maryland, Washington, D. C, Ohio, Kentucky,
Missouri, Texas and Tennessee.

Thionia eUiptica (Germar), 1830

(Plates L. LI, LII, LIII, LV, LVI)

Germar. Thons. Ent. Arch. II, p. 51 Ussus).

Dozier, Herbert L. The Fulgoridae or Plant-Hoppers of Mississippi, Including Tliose
of Possible Occurrence. Miss. Agric. Exp. Sta. A. & M. College, p. 105, 1928.

Comparative notes. This species measures 7 mm. to 7.5 mm.
from apex of head to the tips of the tegmina. It is distinguished
from other species in the genus by the greater proportional width of
the tegmina, the deeply concave basal margin of the frons, by hav-
ing only one longitudinal carina on the frons which extends down
the median line and is sliarply elevated and by the very deeply con-
cave vertex with its greatly elevated anterior corners.

The general color is a yellowish or greenish-tan, speckled with
brown. In addition, the clavus of each tegmen through its middle
third bears a triangle of dark brown and has a dark apex.

460 The University Science Bulletin

Male genitalia. The tenth abdominal segment or anal flap is
much broader than that of simplex, being about twice as wide as
long. At base it is a broad, flat tube that has its ventral margin
greatly extended caudad into an elongate spatulate lobe which is
slightly emarginate at apex. The eleventh segment shows beyond
the dorsal margin of the tubular part of the tenth segment as a
short, ringlike segment, bearing a short, blunt stylus.

Externally the harpagones (genital styli of authors) appear on
the ventral side of the body as two broad, triangular plates which
closely adjoin along their mesal margins. As viewed from a flat-
tened, lateral view each harpago is a broad, pear-shaped plate, nar-
rowed at base, over again as wide across apex and with its dorso-
apical angle extended dorsad into a long process, bearing a pos-
teriorly bent external flap at its base. (See drawing 7, plate LVI.)

The aedeagal structure is a bilaterally symmetrical, broad, tubu-
lar organ with the aedeagus and theca about equally sclerotized and
therefore difficult to distinguish. The theca covers the aedeagus as
a cylinder for about half its length, then splits on each side into a
dorsal flap which extends caudad, completely covering the aedeagus
to the tip, and on its ventral side forms a rounded caudal projecting
lobe just posterior to the aedeagal hook. On the ventral side the
theca extends almost to the tip of the aedeagus, but has each latero-
posterior corner extended still further caudad into a roundingly
pointed mesad curving projection. The aedeagus extends between
the dorsal and ventral flaps of the theca and ends as a broad bilobed
flap which shows between the curved ventral hooks of the theca.
Attached somewhere near its base and extending externally between
the flaps of the theca is a strongly sclerotized hook which abruptly,
bends cephalad and extends as far back as the base of the aedeagus.
About two thirds of the way from the apex this aedeagal hook bears
on its lateral margin a slender, slightly pointed hook.

Location of type. According to Melichar in the Museum in Bud-
apest.

Notes on distribution. Dozier (1928) states that "this is a rare
species, never collected anywhere in numbers." The type locality
is Kentucky. In the Snow Entomological Museum is one specimen
collected at Polk county, Arkansas, by Jack Beamer. In the Na-
tional Museum collection are five specimens collected at Concan,
Texas, by Mr. P. W. Oman.

VanDuzee's catalogue lists it in addition from New Jersey, Wash-
ington, D. C, North Carolina, Georgia, Kentucky and Missouri.

Doering: The Subfamily Issinae 461

Thionia omani n. sp.

(Plates LI, LID
ORIGINAL DESCRIPTION

Size. Length of body from apex of head to tip of tegmen, 7.2 mm.
for the male. Length of tegmen, 6.2 mm.; width of tegmen through
greatest width, 2.2 mm.

Color. General color golden-bronze. Vertex with the extreme
outer margins dark brown, just inside of which is a narrow, light
border, disk mottled brown with a light mesal spot anterior to
center. Pronotum light brown, a yellow longitudinal stripe through
middle of the depressed center and a dark-brown, small depressed
spot on each side of this, each lateral half with about twelve light-
yellow, uniformly sized pimples ; mesonotum reddish-tan through
middle, each lateral carina light yellow, laterad of which is a large
brown spot, lighter at extreme sides and apex. Frons red-brown,
extreme outside margins dark brown, median and lateral carina
and many uniformly round spots and pimples, all light yellow.
Clypeus reddish-tan, with basal or posterior margins and a broad
median longitudinal band lighter tan. Eyes spotted in shades of
brown. Venter of thorax yellow or reddish-tan. Sterna of ab-
dominal segments dark brown through middle, light at the sides.
Tegmina amber, translucent, infuscated in irregular spots or some
small areas near apex light yellow; longitudinal and most of the
cross veins deep tan, a few cross veins near apex light yellow.

Structural details. This is a broad-headed species with the apex
of tegmina greatly narrowed and deflexed. Vertex broad, twice as
wide as one lateral margin, its anterior margin triangularly produced
at middle, its posterior margin triangularly emarginate, the baso-
lateral corners of the frons distinctly showing from above. Pro-
notum roundingly produced anteriorly and shallowly emarginate
posteriorly, its total width approximately six times its total length,
greatly reduced in length behind eyes, due to the great overlapping
of the head and eye region over the pronotum, the latter at this
point being much grooved to allow for the insertion of the eye; all
margins of pronotum being slightly raised, the disk through middle
depressed and in exact center a pair of round, depressed spots.
Mesonotum deeply depressed through middle, a faint median carina
present and the two lateral carinae prominent. Frons a little over
one third longer than basal or posterior margin, which is concave,
the lateral margins slightly diverging at base of apical third, after

462 The University Science Bulletin

which point they extend apically and raesad in the form of a prom-
inent thick carina, the space between the two ends of these margins
being in width about half that of the basal margin ; a median carina
present on the frons, but abruptly ended half way of the length,
two conspicuous lateral carinae also present, which start basally
at the median carina, bow outwardly and end before apex of frons;
the central disk of the frons thickly speckled with round, light-
yellow spots, each lateral disk also speckled in yellow, but these
spots usually more pimple-like. Clypeus approximating the lateral
margins of the frons in length. Tibiae and femora of legs deeply
sulcate. Tegmina very broad at base, prominently buUate at ap-
proximate apex of their basal fourth, from which point they strongly
taper to an angulate apex and from about the middle of the wing,
on are distinctly reflexed; longitudinal veins and a few cross veins
prominent, wing venation characteristic of the genus. Hind wings
characteristic of the genus, being long, with vannal area large, and
with a distinct notch in the prevannal area.

Comparative notes. This species is not easily confused with any
other North American species. It is separated from naso, producta,
quinquata, and hullata, by its broad vertex. From simplex it is
separated by its much larger size and deeper coloring, by the angu-
late margin of its vertex instead of the rounded one of simplex and
by its distinctly bullate and reflexed tegmina. From elliptica it is
distinguished by lacking the strongly concave vertex with elevated
lateral margins as in that species and by having more bulLate
tegmina, at which point they are widest, beyond which point they
fire greatly narrowed and reflexed at apex, while in elliptica the
greatest width of the tegmen is through the apical region, and the
margins are not bent under.

This species probably is more closely related to Thionia variegata
Stal. The writer is not familiar with this species, but judging from
Fowler's illustration of this species and his description, variegata
is unicolorous, brownish-testacesus and does not possess the greatly
narrowed, reflexed tegmina of omani.

The single specimen was sent to Mr. China of the British Museum,
who compared it with Fowler's species of Thionia and Colpoptera
material. He found that it did not compare with these and believed
it to be a new species of Thionia belonging to the variegata group
of Stal.

Location of type. This species was described from one male
holotype, taken in the Organ mountains, Dona Ana, New Mexico,

Doerixg: The Subfamily Issinae 463

August, 1915, collector unknown. The type is in the National
Museum Collection, Washington, D. C.

The species was named in honor of Mr. Paul Oman, of the Na-
tional Museum, who was the first person to recognize this as being
a possible new species and who has done much collecting in this
group of insects, as well as given much assistance to the writer in
compiling this paper.

Thionia naso Fowler, 1905

(Plates LI, LII, LIII, LV, LVI)
Fowler, W. W. Fulgoridae, Biologia Central! Americana. Hoinoptera I, pp. 112-119,

1904.

author's description

Size. Female, length of body from apex of head to tip of tegmen,
7.6 mm. to 8.4 mm.; length of tegmen, 5.6 mm.; width of tegmen,
2.4 mm. Male, length of body, 6.6 mm. to 7.2 mm.; length of teg-
men, 4.8 mm.; width of tegmen, 2.3 mm.

Color. Resembling producta in general color pattern. Dull tes-
taceous except most of ventral surface greenish-tan. Vertex with
thin outer margins dark brown and on each lateral half a brownish
longitudinal band which becomes darker at base, rest covered with
dark-brown specks. Pronotum with a broad, median pale vitta, on
either side of which it becomes darker; slender edges of the pro-
notum as well as some uniformly round, conspicuous dots, dark
brown. Scutellum pale with three irregular brown spots on each
lateral third. Frons yellowish-tan covered with prominent dark-
brown spots; the basal angles and carinae, especially at base, black.
Postclypeus for the most part dark brown with a conspicuous light
vitta down center and a lesser one on each side. Genae, except at
base and apex, ivory. Pleural flap of the pronotum with a light
triangle on lower edge, which sends a slender wedge toward the eye.
Tegmina considerably variegated and quite variable ; the slender 2d
A cell which extends along the entire claval margin, dark through
middle; first anal cell conspicuously but irregularly clouded with
dark; rest of tegmina with dark veins clouded with dark patches
here and there; sternal regions light, marked with dark through mid-
dle and at the sides. Legs light, profusely covered by conspicuous
dark-brown spots, some qf which blend with others in places.

Structural characteristics. The vertex much longer than broad
and greatly produced cephalad beyond the eyes; the basal portion
nearly square, the apical portion extended forward in an acute angle
to such an extent that one edge of the angle is equal to the lateral

464 The University Science Bulletin

edge of the basal region, making all five edges of the vertex approxi-
mately equal; sides carinate and an abbreviated carina on median
line. Frons, elongate, slightly parallel through middle; basal and
apical widths about equal; three longitudinal carinae present, con-
verging at base and extending approximately four fifths of the
length ; posterior margin deeply emarginate for the reception of the
clypeus. Clypeus elongate, strongly convex. Pronotum at middle
about one half the length of the vertex, strongly advanced and ob-
tusely angled between the eyes, truncated behind, ecarinate, with
two impressed points on the disk. Mesonotum slightly longer than
pronotum, its sides arcuated, a lateral carina faintly visible on
either side. Tegmina slender, and costal margin angled as in pro-
ducta, distinctly inflated at a point corresponding to apex of basal
fourth. Venation characteristic of this genus, the longitudinal veins
conspicuously elevated, cross veins few and indistinct.

Male genitalia. Since the general features of the male genitalia
correspond to those in the species producta the reader is referred to
the description of these structures under that heading in the dis-
cussion of that species.

Comparative notes. Since this species more closely resembles T.
producta, the comparative notes given under this same heading in the
description of that species will apply to vaso. As is pointed out
there, these two species can readily be distinguished externally.
However, the matter is somewhat complicated when it comes to
separating these two species by the male genitalia, for it is apparent
that there is not much difference in these usually valuable char-
acters. For this reason some writers might believe that it would be
better to call one a variety than to leave them as two distinct species.
The writer, however, has done the latter thing instead, mainly be-
cause of the idea that taxonomy in a large measure is a matter of
convenience, and since the two are so easily separated by other
characters it would seem less confusing to separate them into two
species.

In the genitalia there are one or two minor differences (see draw-
ings) which can be noted. The dorsoapical processes of the theca
seem less pronounced in omani than in producta. Secondly, the
apices of the aedeagal hooks in producta are always less sclerotized
and seem more slender and tapering, so that at first glance they are
easily not noticed. In omani these apices are more heavily sclero-
tized and seem somewhat bulbous at the end.

Location of types. Fowler described this species from one speci-

Doering: The Subfamily Issinae 465

men collected by Dr. E. D. Ball at Jalapa, Mexico. Doctor Ball
has this type now in his private collection. He has very kindly
compared a male and female specimen with the type. These two
specimens and about thirteen others have been taken at Concan,
Texas, by Mr. Paul Oman of the United States Museum staff.

Notes on distribution. Doctor Ball wrote the following notation
to the author after comparing the Concan, Texas, material with the
type. "In reference to the species, it is typical Thionia 72aso Fowler
as compared with the type and with material which I have from the
Santa Rita, Huachuca and Chiricahua mountains."

Thionia producta Van Duzee, 1908

(Plates LI, LII, LIII, LV, LVI)
Van Duzee, E. P. Studies in North American Fulgoridae. Proe. of the Academy of
Natural Sciences of Phil. LIX : 494, 1908.

Comparative notes. This species is easily distinguished from
simplex, elliptica, bidlata, and quinquata by the vertex, which is
angulately produced cephalad for a considerable distance beyond
the eyes. The basal portion is almost square, with the lateral
margins parallel and slightly elevated as thin edges and the posterior
margin shallowly and roundingly emarginate. The apical portion
of the vertex, starting at the eyes, is produced cephalad in an angle,
a little less than a right angle and each edge of which is about half
the length of the lateral margin. The frons in this species is
elongate, being not quite twice as long as wide and bearing three
longitudinal carinae which are united at base, but which extend
only approximately one half to two thirds of its length. The
tegmina are narrower than in bidlata and have the costal margin
more angled.

T. producta more closely resembles T. naso Fowler than any
others in the genus. From this species it is separated externally by
its smaller size and the shapes of the vertex and frons. In naso
the vertex is similar to that of producta at base, but has the apical
region angulately produced cephalad to such an extent that one
edge of the angle is much longer than either of the lateral margins
or the length of the eye, while in producta this line is shorter than a
lateral line or the length of the eye. The frons in naso is proportion-
ally more slender, being over twice as long as wide and its width at
base where the vertex and frons meet in an angle is approximately
equal to that across its extreme apical margin, while in producta
the basal width at this same point is less than the width at apex.

30—4141

466 The University Science Bulletin

Notes on variation. This species seems very variable in color.
In the original description much black was noted on irons, vertex,
pronotum, and a broad vitta on the tegmina following the inner
margins of the scutellum. In many specimens this black is greatly
reduced or lacking in many places.

Structurally a variation is noted in the extent that the vertex is
produced cephalad. A series of specimens show the vertex longer
than in the typical forms, but not nearly as long as in naso. The
writer places these longer-headed forms with producta, although
Doctor Ball stated in a notation concerning naso Fowler that he
had material of this species from the Santa Rita mountains, Arizona.

Male genitalia. The anal flap (tenth abdominal segment) in this
species is so elongate that its apex bends caudad, snugly covering
the extreme apices of the genital styli on the dorsal surface. Its
length is a little longer than three times its width. At base the
tenth segment is a flattened tube with parallel lateral margins. Its
ventral margin extends. caudad as the parallel-sided flap, which is
truncate and slightly tapering at apex and not quite twice as long as
the basal part. The eleventh segment is visible beyond its dorsal
margin as a narrow rihglike segment bearing a moderately long
stylus.

The harpagones (genital styli) are visible externally as broad,
triangular plates whose inner margins are closely adjoined. From a
flattened lateral view each harpago is a broad, pear-shaped plate,
narrowed considerably at base, where it is attached to the body by
a slender stalk. Its apex is considerably widened and its dorso-
posterior region bears a foot-shaped flap which extends first caudad,
then suddenly bends dorsad, where it projects dorsad as a roundingly
pointed projection.

The aedeagal structure is similar to that of other species, especially
that of bullata. It is bilaterally symmetrical. On the ventral side
its posterior margin is truncate through the middle, but at the sides
is deeply notched to allow the aedeagal hooks to appear externally
as anteriorly projecting, well-sclerotized hooks. Laterad of the
notches the theca projects caudad on either side as slender, some-
what pointed processes. From a lateral view it appears much longer,
covering practically the entire aedeagus. Its extreme dorsal and
apical region is modified into two projections, the anterior one be-
ing a rounding, cephalad-projecting lobe and the posterior one being
a short, pointed, caudad-projecting hook. The aedeagus appears
only at the extreme apex, where it appears as a broad, elongate lobe,
emerging from between the flaps of the theca. It bears on each side

Doering: The Subfamily Issinae 467

a long, narrow, well-sclerotized process which emerges from the
notch of the theca, extends directly ventrad a short distance and
then bends abruptly forward, reaching not quite to the base of the
theca. Each aedeagal hook bears on its outer margin at approxi-
mately the base of the apical fourth a small external spine. In situ
these hooks are crossed on the ventral side.

Notes on distribution. Van Duzee described this species from one
pair taken at Rifle, Colo., in July. Mr. Van Duzee was kind enough
to send to the writer a metatype male taken in the Huachuca moun-
tains, Arizona, in August. Mr. Paul Oman has taken a series of
fourteen at Concan, Texas, in June. In the Snow Entomological
Museum at the University of Kansas is a large series taken in Ari-
zona at the following places; Huachuca mountains, Santa Rita
mountains, Yavapai, Prescott, Jerome, Miami, Coconimo county,
Chircahua mountains and Oak Creek canyon; and in Utah from
Cove Fort, Cedar City, Monroe and Salina. In 1936 Dr. R. H.
Beamer collected a series from Leakey, Texas, on July 8.

Location of types. In the collection of Dr. E. P. Van Duzee, San
Francisco, Cal.

Host plants. Collected by Dr. R. H. Beamer from cedar (1936).

Thionia qidnquata Metcalf, 1923

(Plates LI, LID

Metcalf, Z. P. A Key to The Fulgoridae of Eastern North America With Descriptions
of New Species. Jour, of the Elisha Mitchell Scientific Society 38; p. 190, 1923.

Comparative notes. Metcalf in his original description states that
this species may be recognized by the shape of the vertex and by
coloration.

The general color is almost uniform light brown, which is almost
uniformly covered with small, dark points, and the veins are also
dark.

In size it ranks as one of the larger species in the genus, as the
female measures at least 8 mm. in length from tip of head to apex
of wings.

The vertex is more distinctly chevronlike in shape than in any
other species in the genus due to the sharper angles, especially the
anterior one, and the fact that the six sides of the chevron are prac-
tically equal ; the lateroanterior angles of the vertex are posterior to
the anterior margin of the eye and, therefore, more of the frons is
visible from above than in such species as hullata. The frons is
broad as in hullata, being not much longer than wide; it bears three
distinctly elevated longitudinal carinae, which are united at base;

468 The University Science Bulletin

its basal margin is only slightly concave. The longitudinal veins of
the tegmina are prominent and elevated; the cross veins are more
conspicuous than in other species.

Note. The writer was in possession of only one specimen for
study. This was a female, and therefore no data in regard to the
male genitalia can be given.

Notes on distribution. Metcalf states that the type female was
collected at Raleigh, N. C, in September. One specimen in the
United States National Museum was taken at Thompson Mills, Ga.,
concerning which no data was given in regard to date or collector.

Thionia bidlata (Say), 1830

(Plates LI, LII. LIII, LV, LVI)
Say, Thomas. Jr. Acad. Nat. Sci. Phila. VI, p. 240; Comp. Writing, II, p. 375. Flata.

Comparative notes. A common species which is rather variable in
color and, therefore, not always easily identified. The usual pat-
tern has a light vertex marked with a dark longitudinal band on
each lateral half (sometimes, however, reduced to a spot), which
usually continues, although sometimes irregularly, across pronotum
and mesonotum; tegmina unicolorous light brown or tan with a
darker indefinite band in the form of a Y which is interrupted
through the middle.

Size from tip of head to apex of tegmina 6.4 mm. to 7.6 mm.

The structural characteristics by which it can best be distinguished
are the shape of the frons, the vertex and the tegmina. The frons is
only slightly longer than wide; its lateral margins are roundingly
and evenly emarginate, making the greatest width through apical
third; it bears three distinctly elevated carinae which are united at
base, and its basal margin is very slightly concave. This species re-
sembles quinquata in many respects. It differs from it by its smaller
size, shape of the vertex and tegmina. In the latter species the
chevronlike shape of the vertex is more pronounced, due to the six
sides of the chevron being approximately equal in length. In
bullata the lateral margins are proportionally longer and this brings
the lateroanterior angles in line with the anterior margin of the
eyes or more commonly anterior to the eyes, while in quinquata this
angle is posterior to the anterior line of the eyes. The tegmina of
quinquata are distinctly inflated at base of apical fourth and only
sligiitly so in bullata.

Male genitalia. Externally the male of this species can be dis-
tinguished very easily by the conspicuous spatulate projections from

Doering: The Subfamily Issinae 469

the anal flap, which extend ventrad around the harpagones to such
an extent that they ahnost converge at middle on the ventral side
of the body. The tenth abdominal segment (anal flap) is con-
spicuously different from other members of the genus studied. At
base it is a broad, flat tube, but apically it has its ventral and lateral
margins projected caudad as a greatly expanded plate, whose latero-
posterior corners are drawn out into long spatulate lobes (see above)
and whose posterior margin through the middle is triangularly pro-
duced. The eleventh abdominal segment shows beyond the dorsal
margin of the tubular part of the tenth segment as a short ringlike
segment which bears an abbreviated stylus.

Externally the harpagones (genital styli of authors) appear on the
ventral side of the body as two broad, triangular plates which closely
adjoin along their mesal margins. As viewed from a flattened,
lateral view, each harpago is a broad, pear-shaped plate, narrowed
considerably at base where it is attached to the body by a slender
stalk, but over twice as wide across the apex as at base and with
a slender footlike flap extending first caudad, then suddenly bending
dorsad as a roundingly pointed flap.

The aedeagal structure is of the siynylex type. It is bilaterally
symmetrical. On the ventral side the theca covers the aedeagus
for three fourths of its length ; it has a truncate posterior margin and
each lateral third is extended laterally as a rounded flaplike lobe
which is just caudad to the aedeagal hook at a point where it
emerges from the notch in the theca. From a side view the theca
shows a modification in the dorsoapical region, namely, by having
two projections, an anterior one which is a rounded lobe that ex-
tends cephalad over the aedeagal structure for about half its length,
and a posterior one which extends caudad and slightly dorsad as a
slender, gently curving hook. The aedeagus shows as a thick lobe
between the processes of the theca and bears two long, narrow,
pointed processes, which protrude, one on each side, from the lateral
notches in the theca and bend directly cephalad, extending as far
forward as the base of the aedeagus. Each of these aedeagal hooks
bears on its outer margin, at approximately the base of the apical
sixth, a small external spine.

Notes on distribution. Say, in the original description, states that
it inhabits the United States.

Specimens were available for study from Alabama, Arkansas,
Florida, Georgia, Louisiana, Mississippi and South Carolina. In
addition to these states, Dozier (1928) gives the following: New

470 The Univeesity Science Bulletin

Jersey, New York, Ohio, Pennsylvania, and Ontario. He states that
both nymphs and adults were taken on oak.

Van Duzee's catalogue adds North Carolina.

Location of type. Do not know.

Thionia ocellata Melichar, 1906

Melichar, Leopold. Monograph des Issiden (Homoptera). Abk. k. k. Zool. Bot. Ges.
Wein III, 1906.

Metcalf, Z. P. The Fulgoridae of Eastem North America. Jour, of Elisha Mitchell
Society, Vol. 38: 158, 1923.

Comparative notes. The writer did not have any specimens of
this species for study. Mr. Paul Oman says that there are no speci-
mens in the National Museum, and that he is not familiar with the
species.

Metcalf (1923 ) , in his key to the species, distinguishes it by having
vertex as long as broad, frons not arched, without a transverse yel-
low band, all reddish or yellowish-brown, vertex rounded anteriorly
and tegmina with an ocellated spot near the apex.

The size of the species is given as 5% mm.

Location of types. In the Wien Museum, Austria.

Notes on distribution. Apparently described from one female, for
which the locality is given as North America. The writer believes
that this species probably does not occur in North America, north of
Mexico.

Thionia transversalis Melichar, 1906

Melichar, Leopold. Monograph des Issiden (Homoptera). Abk. k. k. Zool. Bot. Ges.
Wien III, 1906.

Metcalf, Z. P. The Fulgoridae of Eastern North America. Jour, of Elisha Mitchell
Society, Vol. 38: 158, 1923.

Comparative notes. The writer did not have any specimens of this
species for study. Mr. Paul Oman says that there are no specimens
in the National Museum and that he is not familiar with the species.

Metcalf (1923) , in his key to the species, distinguishes it by having
vertex as long as broad, frons slightly arched and with bright yellow
transverse band.

The size of the species is given as 8 mm.

Location of type. Melichar states that the type female is in the
museum in Wien, Austria.

Notes on dutribution. The species was evidently described from
one female, for which specimen the type locality is given as North
America. The present writer is of the opinion that this is not found
in North America, north of Mexico.

Doering: The Subfamily Issinae 471

The Genus Picumna Stal

Stal, Carolus. Hemiptera Mexicana — Enumeravit Speciesque Novas Descripsit. Stet. Ent.
Zeit. XXV: p. 53.

Fowler, \V. W. Fulgoridae. Biologia Centrali Americana. Homp. I; 113-119, 1904.

Melichar, Leopold. Monograph der Issiden (Homoptera.) Abli. k. k. Zoiil. Hot. Ges.
Wien III, 190C.

Comparative notes. In size varying from 6-7 mm. long. General
color, brown, marked witii testaceous and yellow.

Some of the distinctive characteristics of this genus arc: the frons
narrowed at base with one median carina distinctly visible and two
lateral carinae faintly indicated or showing only at the extreme
basal end; the clypeus keeled through the middle; vertex concave
across disk; pronotum short with very abbreviated lateral areas,
anteriorly curved and posteriorly straight; scutellum with three
parallel longitudinal keels, the median one not always distinct;
tegulae conspicuous; tegmina elongate, oval, weakly produced at
sides and with thick longitudinal veins, the cross veins lacking or
scarce, clavus long, extending to base of apical fourth of tegmen;
hind wings long, deeply notched at margin; hind tibia with usually
four lateral spines (rarely five) and a crown of spines at apex.

The wing venation in this genus shows the following characteris-
tics: vein Sc is unbranched and, therefore, there is no expanded
costal area; Sc, and R are united for only a short distance at base;
M is always branched into two branches, usually about center of
tegmen; vein Cu^ is branched always posteriorly to the branching
of M, but in various specimens varying in its distance from apex of
tegmen; Cu^ follows the anal suture; 1st A and 2d A uniting some
distance from apex of clavus.

This genus resembles the genus Thionia rather closely, but is
easily distinguished, as Stal points out, by its quadrispinose hind
tibia and vein Cu^ being branched; in Thionia the hind tibiae are
bispinose and vein Cu^ is simple.

Key to Species

la. Dark brown, mottled with yellow ; costal margins of tegmina as viewed froin
above not parallel, somewhat expanded at base so that greatest width of
tegmen is in line with apex of scutellum Picumna chinai n. sp., p. 473

lb. Dark brown, mottled with yellow; costal margins parallel or slightly rounding,
not expanded at base, greatest width of tegmina throuih middle.

Picumna macuiatn (Melicliar), p. 471

Picumna maculata (Melichar)

(Plates II, IV, V. VI, VIII)

Issomorphus maculatus Mel. Melichar, Leopold. Monograph der Issiden, Homoptera.
Abh. k. k. Zool. Bot. Ges. Wien III, 1906.

Issomorphus maculataus Mel. Met calf, Z. P. The Fulgoridae of Eastern North America.
Jour, of the Elisha Mitchell Society XXXVIII, 1923.

Picumna ovatipennis Walker. Fowler, W. W. Biologia Centrali Americana. Homoptera
I; 113-119, 1904. Figures a variety, fig. 30, plate 12, which may be maculata Melichar.

472 The University Science Bulletin

Comparative notes. Issomorphus maculatus in this paper is
placed under the genus Picumna because it has all the characteristics
of this genus. In appearance it is very similar to other species in
the genus. Melichar in his original description of the genus Issomor-
phus mentions the presence of five spines on the hind tibia which he
says separates it easily from the European genus Issus, and which
would also separate it from any North American genus. The present
writer has been unable to locate five spines on any specimen, and
believes that Melichar must have counted the large lateral spines
which make up part of the apical crown or circlet of spines.

This species in size and shape resembles very closely two Mexican
species, Picumna ovatipennis Walker and Picumna varians Stal. In
many collections the former name has been given to this species. It
also resembles in color pattern and size Picumna chinai n. sp. For
comparison of these species see the notes under this heading in the
description of the latter.

A color variation occurs in this species, some specimens being
much more mottled than others and some with uniform brown
tcgmina, except for a large yellow spot along the costal margin.

Male genitalia. The tenth abdominal segment (anal tube or flap)
is a broad, flat tube at base that has its ventral margin extended
caudad as a broad, spatulate truncate lobe which is wider across
extreme apex. The eleventh segment shows beyond the dorsal
margin of the tenth segment as a short, ringlike segment and bears
a long, slender, pointed stylus.

Externally the harpagones (genital styli of authors) appear on
the ventral side of the body as two triangular, pointed plates which
are closely adjoined along their mesal margins. As viewed from a
flattened lateral view each harpago is a broad pear-shaped plate,
narrowed at base, greatly broadened out across apex with the dorsal,
apical angle extended dorsad into an elongate process which is
slightly pointed at its extreme apex and at base bears an ex-
ternal, liplike process.

The aedeagus and theca are difficult to distinguish. Together they
appear as a bilaterally symmetrical, broad, tubular structure. From
a lateral view the theca shows its apical margin broadly notched
through middle, due to its ventral region being greatly extended
caudad as a rounded lobe which bears on each side a rounded liplike
process just posterior to each aedeagal hook, and its dorsal region
extending caudad as three distinct processes, the one through mid-
dle being a broad semimembranous flap and the two on each side

Doering: The Subfamily Issinae 473

being slender, sinuately curved, hooklike processes which are sharply
pointed at apex. The apical region only of the aedeagus shows.
This part consists of a broad, bilobed flap extending between the
dorsal and ventral flaps of the theca, and two sclerotized hooks, one
on each side, which protrude from the notches of the theca and im-
mediately bend caudad for a short distance, then make a right-
angled bend which causes them to extend cephalad over the aedeagus
almost to its base. Each aedeagal hook, as viewed from the ventral
side (drawing 3, plate LIV), is slender and cylindrical at base, then
suddenly constricts to a narrow rod and then flattens out as a spatu-
late flap, bearing on its mesocephalic corner a slender, recurved hook.

Notes on distribution. Melichar lists Las Vegas, Hot Springs,
New Mexico, as the habitat of this species. In the Snow Entomo-
logical Collection at the University of Kansas are many specimens
taken by R. H. Beamer from the following places in Arizona:
Coconimo county, Yavapai county. Oak Creek canyon, Grand Can-
yon, Huachuca mountains, Santa Rita mountains, the Chircahua
National Monument, and Granite Dell. One specimen was collected
by Mr. Beamer from Luna, N. Mex., in July.

Location of types. In the National Museum at Washington, D. C,
are three syntypes of the species Issoinorphus maculatus Melichar.
Mr. Paul Oman has kindly made comparisons of these types and
finds that two of them, although somewhat broken, fit the original
description and figures made by Melichar for this species. One of
these we are designating as the lectotype of the species. The data
for this type is "Las Vegas, H. S., N. M., Aug. 13. Barber and
Schwarz, collectors." According to Mr. Barber the H. S. means Hot
Springs, which are at the base of the hills about six miles from the
town of Las Vegas and in the edge of the pinon pine belt.

The third specimen of the group is the new species, Picumna chinai,
which follows this description.

Host plants. Dr. R. H. Beamer collected a series of this species at
Silver City, N. Mex., from pine (1936).

Picumna chinai n. sp.

(Plates L, LU, LIII, LIV, LVI)

Picumna ovatipennis Walk. Fowler, W. W. Biologia Central! Americana. Homoptera
I, pp. 113-119, 1904. (Probably is his figure 29, plate 12.)

Picumna ovatipennis Walk, ^etcalf, Z. P. The Fulgoridae of Eastern North America.
Jour, of the Elisha Mitchell Society XXXVIII, 1923.

ORIGINAL DESCRIPTION

Size. Length of body from apex of head to tip of tegmen 5.2 mm.
to 5.6 mm. Length of tegmen 4 mm. to 4.4 mm.; greatest width of

474 The University Science Bulletin

tegmen 1.6 to 1.8 mm. This species has the size and appearance of
maculata (Mel.).

Color. This species has the typical mottled color pattern in black-
ish brown to cream that is characteristic of the genus. Vertex, pitch-
brown except for a median light tan longitudinal stripe and a spot
of same color in center of each lateral half. Eyes, light brown or
tan. Pronotum, pitch-brown except a median yellow stripe and the
lateral carinae somewhat lighter. ]\Iesonotum, light tan, mottled
with light brown and all margins and carinae lighter. Frons, light
tan, traversed by irregular tracings of dark brown which usually
becomes a solid patch across basal or posterior margin and roundish
spots on apical lateral corners, carinae brown. Clypeus, yellow,
dark brown at sides, each lateral half bearing 6 to 7 oblique brown
bands. Underside of thorax yellow with conspicuous dark-brown
spots on each sclerite. Coxae of forelegs, light yellow, with a con-
spicuous brown spot on anterior half; femora of legs striped lon-
gitudinally in yellow and brown; tibiae reddish-brown; tarsal seg-
ments light, shading to reddish-brown on margins and into black-
tipped claws. Abdominal segments of male from underside light
yellow, harpagones infuscated. Abdominal segments of female from
underside light at sides and margins, darker through middle; ovi-
positor valves pitch-brown. Tegmina, pitch-brown, irregularly
mottled and speckled with light, a large semihy aline light spot on
costal margin approximately at middle, wiiich is somewhat triangu-
lar in shape, with the apex of the triangle directed mesad.

Structural details. Vertex transverse, concave, lateral margins
slightly diverging posteriorly, anterior margin evenly rounding, but
not angulate as in maculata, posterior margin deeply triangularly
emarginate. Pronotum short, broader than head, slightly sunken
through middle, shorter behind eyes and the anterior part of each
lateral arm greatly sunken to allow for the overlapping of the eye
onto it, behind which it is elevated into a sharp carina. Mesonotum
triangular, not much broader than long, only slightly sunken through
middle, median carina only faintly indicated, a lateral carina on
each side conspicuous. Tegulae very large. Frons elongate, its
lateral margins diverging apically so that at apex it is not quite a
third wider than across the basal margin; the basal margin deeply
concave; a median carina and a lateral carina on each side starting
at the same place, slightly below the basal margin, the two lateral
ones outwardly bowing and then running parallel with median one.
Clypeus almost as broad at base as the frons, its greatest length
about equal to one lateral margin of the frons. Tegmina held hori-

Doering: The Subfamily Issinae 475

zontally on body, their claval and costal margins subparallel, their
greatest width just anterior to middle, where they are considerably
flared as viewed from above.

Male genitalia. Anal flap (10th abdominal segment) forms a
characteristic flattened tube at base, the ventral margin of which is
then extended into an elongate, parallel-sided, truncate lobe. The
eleventh segment shows slightly beyond the dorsal rim and bears a
slender, fingerlike stylus, which is many times longer than wide.

The harpago is extremely broad in this species, being almost tri-
angular in shape. It is a flat lobe, slightly inflated through middle
and with its extreme dorsoposterior corner extended cephalad into a
somewhat bluntly tapering projection. At the base of this projec-
tion is a flat, external ventrad-projecting flap.

The aedeagal structure is bilateral and of the same general shape
of macidata, but broader. From a lateral view the theca shows its
apical margin broadly notched through middle, due to its ventral
region being greatly expanded caudad as a rounded lobe, which
bears on each side a rounded liplike process just posterior to each
aedeagal hook, and its dorsal region extending caudad as three dis-
tinct processes, the one through middle being a broad semimem-
branous flap and the two on each side being slender, twice angled
hooks which end in a sharp apex. The apical region only of the
aedeagus shows. This part consists of a broad, bilobed flap extend-
ing between the dorsal and ventral flaps of the theca and two sclero-
tized processes, one on each side, which protrude from the notches
of the theca, immediately bend caudad, then make a right-angled
bend, which causes them to lie over the aedeagus, where they reach
to almost the base of the latter. Each aedeagal hook as viewed from
the ventral side is slender and cylindrical at base, then suddenly be-
comes slightly constricted, after which it suddenly expands, becom-
ing a spatulate lobe with its mesocephalic angle forming a small,
recurved hook.

Comparative notes. In collections of the Issidae taken in North
America, north of Mexico, there are tw^o species of this group rep-
resented in fairly good numbers. These two species have been de-
termined by authors as Issoin&rphus maculatus Melichar and
Picumna ovatipennis Walker, and have been indiscriminately and
interchangeably classified as such. In order to straighten out which
was which the writer sent specimens from each species to Mr. China,
of the British Museum, to compare with the type of Picumna ovati-
pennis Walker. Mr. China reported that neither of these United

476 The University Science Bulletin

States forms were ovatipennis Walker, and sent a homotype (com-
pared by him) of the latter to the writer for study. This homotype
is described in the following description and figures are given for it.
Mr. China suggested that the ovate United States species was 7s-
somor'phus 7naculatus Melichar and the other one, with the broad-
ened tegmina at base, a new species.

Following Mr. China's comparison the writer then asked Mr.
Oman, of the United States Museum, to again check over the type
specimens. Mr. Oman reported that two of the three type speci-
mens of Issomorphus maculatus Melichar were the ovate species and
one was the broad-winged one. Because the ovate one fits Melichar's
figure better we have designated it as the lectotype, and since it un-
doubtedly is Picumna we have changed the name to Picumna macu-
lata (Melichar). See preceding description. The broad-winged
form is herein described as new, with the name Picumna chinai.

The species of the genus Picumna resemble each other most closely
in coloration and size. In these two respects P. maculata (Mel.)
and P. chinai n. sp. are almost indistinguishable. Yet, these two are
easily separated by the shape of the tegmina. P. chinai, as viewed
from above, is a broad, flat species, more the shape of Ulixes scutatus,
due to the great expansion of the wings at the base which makes the
tegmina broadest at a point in line with apex of scutellum. P. macu-
lata (Melichar) has the characteristic shape of the genus, which is
ovate, as viewed from above, with the greatest width of the tegmina
through the middle. Tlie frons is slightly more narrowed at base
and the clypeus slightly broader in chinai than in maculata. The
genitalia of these two species are similar and yet show distinctive dif-
ferences, which can best be seen by studying the drawings.

The Mexican species, Picumna ovatipennis Walker, is easily dis-
tinguished from the North American species occurring in the United
States, by the shape of the vertex and frons. Although Walker de-
scribed the vertex as transverse in this species, it is subequal in
length and width, which gives it a much longer appearance. In
P. maculata (Mel.) and P. chinai n. sp. the vertex is at least one
fourth wider than long and has the appearance of being quite broad
and short. The frons of P. ovatipennis is greatly narrowed at base
and the clypeus much shorter than either of the other two species.

The writer has not seen any of the other Mexican species which
might be confused with any of the above three. Mr. China believes,
as does the writer, that both Fowler and Melichar thoroughly mud-
dled up the genus Picumna. He states that P. I'arians Stal, as de-

Doering: The Subfamily Issinae 477

termined by Fowler, seems to be closely allied to ovatipennis Walker.
This idea is borne out by the fact that in Stal's description of
varians he states that the vertex is equal in length and width.

Notes on distribution and types. The species was described from
numerous examples taken at various points in Arizona. The species
is named after Mr. China, of the British Museum, who has so kindly
assisted the author in the identification of numerous species and who
first suggested that this was a new species.

Holotype male and allotype female collected in the Santa Rita
mountains of Arizona, on August 18, 1935, by R. H. Beamer. Para-
types from the following places: 14 females and 9 males from the
Santa Rita mountains; 16 females and 11 males from the Huachuca
mountains; 2 females and 2 males from the Chircahua mountains;
2 females and 2 males from Patagonia; one female from St. Catalina
mountains, and two males from the Dragoon mountains.

Holotype and allotype, and the majority of the paratypes in the
Snow Entomological Collection at the University of Kansas.

Picumna ovatipennis (Walker)

(Plate L)

Walker, Francis. List of the specimens of Homopterous Insects in the Collection of the
British Museum. Supplement, p. 88, 1858, as (Issus).

Size. Length of body, 4.8 mm. Length of tegmen, 4.5 mm. ; width
of tegmen, 1.9 mm.

Color. General color greenish-tan, marked with brown. Vertex
yellow through middle, lateral margins greenish, apical third and a
spot in each lateroposterior corner brown. Frons, yellow-green,
marked with dark in the following places; basal angles dorsad of
carinae dark brown, lateral carinae faintly indicated almost to apex
as brown lines which fade out apically, disk at base between lateral
arms of carinae reddish-brown, followed by an irregular transverse
band of light, another dark band through middle followed by a light
area at apex; in each lateral disk through middle is a vertical row
of about six round, brown spots, the expanded apical corner is
mottled in brown. Clypeus greenish-j^ellow with six oblique bands
indicated on either side, apical half dark brown. Genae yellow ex-
cept a brown crescent around base of each antenna. Antenna dark
brown with lighter brown sense organs. Pronotum greenish-yellow
with two brown spots in apex of triangle formed by the lateral
carinae and a brown spot at each extreme lateral edge. Pleural
parts below yellow margined, rest dark brown except a light streak
through middle. Rest of thorax below yellowish-green, splotched

478 The University Science Bulletin

with brown, from above light brown. Abdominal segments, from
above light brown, at sides considerably darker and extreme pos-
terior margins light. From below abdominal segments brown
through middle, mottled at sides, extreme posterior margins green-
ish-yellow. Ovipositor dusky brown. Legs, femora brown with a
broad transverse light band through middle; tibiae speckled; tarsal
segments reddish-brown, somewhat darker on apices, tarsal claws
dark brown. Tegmina yellowish-hyaline marked with reddish-
brown in the following places : a broad triangle through middle with
its apex ending on claval suture at about middle of wing, an oval
spot on extreme apical costal angle, another large rectangular area
between this spot and apex of corium, smaller spots scattered over
other parts of corium, clavus with a larger, irregular oblique band
across middle, a smaller irregular band posterior to this and the
extreme basal posterior area dark brown. Wings pale, transparent
with thick brown veins.

Structural details. Vertex with length and width equal, consider-
ably concave through disk, making it appear narrower than it is,
its anterior margins roundingly converging to a sharp pointed apex
from above, thus allowing considerable of frons to show on either
side. Width of eyes considerably greater than width of vertex.
Pronotum considerably wider than head, distinctly triangulate an-
teriorly with its two lateral carinae converging to a sharp point as
they touch the anterior margin. Mesonotum large, slightly over
twice the length of the pronotum at middle, a transverse crease fol-
lowing along the anterior margin and ending on each side at the
lateral carina, no median carina visible, the central disk somewhat
depressed. Abdominal segments collapsed in the dried specimen.
Anal flap (10th segment) long, the extended flaplike part about one
third longer than the tubular part. Genital stylus of the eleventh
segment, a long, slender, fingerlike projection. Tegulae conspicuous.
Tegmina subparallel, margined with the costal margin only slightly
rounding, apex uniformly rounding, and its greatest width, if any,
in line with angle of the claval margin. Hind wings large, with its
three areas at apex about equal in width.

Wing venation same as for the genus in general characteristics. In
particular, M branches into two branches considerably anterior to
middle, then each branch again divides in the apical region. Cu^
branches only a slight distance posterior to middle ; anal veins unite
at about base of apical fourth of clavus.

Comparative notes. This Mexican species apparently has not
been collected north of Mexico at the present time. Two or three

Doering: The Subfamily Issinae 479

northern species have been wrongly determined as this species in
collections and in the literature. For comparative characters to sep-
arate the several species see the notes under this heading in the de-
scription of Picumna chinai n. sp., just preceding.

Mr. China very kindly loaned the writer a female homotype com-
pared by him with the type. Drawings of this specimen are found on
plate L, figs. 1, 2, and 3. In studying this type specimen with Walk-
er's description the present writer finds the following inconsistencies
in the description: Walker states that the vertex is transverse,
whereas, in the specimen it is subeciual in length and width ; that the
abdomen is black — in this specimen it was testaceous brown; that
the forewings are black and tawny towards the base — in this speci-
men they are tawny with brown markings; that the forewings have
a tawny mark on the anterior border — this specimen had a large
costal spot which also continued transversely across wing; that hind
wings are grayish vitreous with black veins, while here they are pale
yellow with light-brown veins.

Location of types. British Museum.

The Genus Euthiscia Van Duzee

Van Duzee, E. P. Expedition California Academy of Science to Gulf of California. Prac.
Cai. Acad, of Sc. XII, p. 193, 1923.

C omparative notes. According to Van Duzee this genus has the
aspect of Mycterodus and is closely allied to Thiscia Stal. Neither
of these genera are North American. It differs from Thiscia by the
very broad, smooth front, much simpler elytral venation and the
rudimentary wings.

The genus is readily distinguished from other North American
genera by spineless hind tibiae, the very large pronotum as compared
to the scutellum and the broad, much abbreviated and somewhat
bullate tegmina which enclose the body and meet below it.

Van Duzee described two species in the genus, of which one species,
Euthiscia tuberculata, has been taken north of Mexico by Dr. E. D.
Ball, at Tucson, Ariz.

Euthiscia tuherculata Van Duzee, 1923

(Plates I, IV, V, VI. VlII)

Van Duzee, E. P. Expedition California Academy of Science to Gulf of Mexico. Proc.
Cal. Acad, of Sc. XII, p. 193, 1923.

Comparative notes. This species is readily distinguished exter-
nally by several characteristics. On the dorsum of the body are six
tuberculate elevations, two on vertex, two on pronotum and two on
the scutellum. The tegmina are unusual because of their shape and

480 The University Science Bulletin

brevity; being so abbreviated at apex that little of the tegmen ex-
tends caudad beyond tip of clavus and their greatest width is at a
point considerably cephalad to their middle. As is typical for the
genus the tegmina are held vertically, are somewhat bullate, and
enclose the body by having their costal margins meet below.

The wing venation shows the following general characteristics:
vein Sc divided, with vein Sc^ (equal costal vein of Metcalf) only
visible a short distance at base; veins Sc,, R, M and Cu all ap-
parently united at base for a short distance; veins Sc2 and R branch-
ing from this main trunk as one vein, but soon dividing into separate
veins; vein M four-branched; vein Cu^ two-branched; vein Cuo
I'unning along claval suture as in other Homoptera ; veins 1st A and
2d A by uniting at apex forming a Y-vein in clavus. All the veins
of the tegmina breaking up into a reticulation at apex so that it is
difficult to trace out the course of the longitudinal veins. Hind
Mdngs lacking.

Male genitalia. Anal flap (lOth abdominal segment) almost three
times as long as broad, narrowed at apex. Eleventh segment ring-
like and with a conspicuous elongate stylus. Both of these segments
almost hidden externally, due to the great compression of the ab-
dominal segments and the dorsal extension of the apices of the
harpagones.

The harpagones in this species are visible externally as two ventral
subrectangular plates meeting at middle. They are proportionally
smaller than in other species in the subfamily. Due to the great
compression of the sterna of the abdominal segments they assume,
also, a more vertical position with their apices extending considerably
dorsad, thus crowding the anal flap and genital stylus backward un-
til the latter touch the abdominal terga and are practically hidden
by the harpagones.

The aedeagus is a broad tubular structure which is bilaterally sym-
metrical. Its apical half is divided into two broad spatulate lobes,
each of wiiich bears four irregular teeth on its laterocaudal margin
and on its ventral side a cephalad-projecting, sharply pointed, elon-
gate flap, which extends cephalad to a point midway the length of
the aedeagus. A broad, three-pronged apodeme is attached to the
base of the aedeagus. A rather complicated theca envelops the
aedeagus. On the ventral side the theca, through the middle, covers
the aedeagus for about two thirds of the length of the latter, but the
extreme lateral angles project caudad much farther and somewhat
dorsad as two sharply pointed processes. On the dorsal side the
theca is more membranous, is depressed through the middle for

Doering: The Subfamily Issinae 481

most of its length and at the apical end projects caudad between the
apical flaps of the aedeagus as a broad, somewhat pointed process.
(See drawing 9, plate LIV.)

Notes on distribution. The species was described from numerous
examples from lower California. Holotype male taken at Los
Angeles Bay and allotype female at Monserrate Island. Types are
m the California Academy of Science Collection at San Francisco.
Dr. E. D. Ball has since taken this species from a gray shrubby
Verbena (Lippia urightii) in Sabino canyon, near Tucson, Ariz.

The Genus Hvsteropterum Amyot and Serville

Amyot and Serville. Hemip. p. 519, 1843.

Fowler, W. \V. Fulgoridae. Biologia Central! Americana. Honioptera I, pp. 113-119,
1904.

Distant, W. L. Fauna of British India, Family Fulgoridae. Rhynchota III, p. 333, 1906.

Melichar, Leopold. Monograph der Issiden (Homoptera). Abh. k. k. Zool. Bot. Gea.
Wien, III, pt. 4, 1906.

Metcalf. Z. P. The Fulgoridae of Eastern North America. Jour, of the Elisha Mitchell
Society XXXVIII, 1923.

Dozier, Herbert L. Fulgoridae. Miss. Agric. Exp. Sta. Bull. 14. 1928.

Comparative notes. The distinguishing characteristics of this
genus are: Tegmina short, either subequal in length and width or
not twice as long as wide, deflected in an oblique or vertical position
to the body, not convex, narrowed and rounded at apex; costal mar-
gin narrow, bent at an angle to rest of wing and lying horizontally
against venter; hind wings small and rudimentary; head not por-
rectly produced in front of eyes; frons centrally and laterally cari-
nate; hind tibia either with one or two stout lateral spines; male
genitalia bilaterally symmetrical.

The general features of the venation are: vein Scj close to costal
margin and from a dorsal view appearing to be the ventral margin
of the tegmen; veins R and Sc^ united a very short distance at base,
each single vein parallel througliout rest of wing; vein M divided
into two branches near middle of wing; vein Cu^ simple; vein Cuj
following the anal suture; veins 1st A and 2d A united near apex
to form the characteristic Y-vein of the clavus. All the longitudinal
veins break uj) into a close reticulation at apex of wing.

Key to Species

1. Larger insects, measuring 6 tu 7 mm. long; dark brown or blackish in color-

ing; reticulate elytra morum Van Duzee, p. 482

Smaller insects, measurmg under b mm. in length ; varying from speckled
dark brown to light cream in coloring 2

2. (1) In lateral view dorsal line of tegmen straight; usually uniformly colored,

marked with numerous dark spots in the cells of the tegmina (occasion-
ally two-toned) 3

31—4141

482 The University Science Bulletin

Somewhat concave through middle; either contrastingly colored in fuscous
and light or cream, spotted with dark in varying degrees 5

3. (2) In dorsal view frons protruding almost equal to length of vertex, lateral

carinae against eyes scarcely elevated; disk of vertex less concave;
anterior margin of pronotum rounded; lateral edge of tegmen cream

colored auroreum Uhler, p. 483

In dorsal view frons protruding not as much as half length of vertex, lateral
carinae against eyes distinctly elevated; disk of vertex concave, anterior
margin of pronotum angulate; tegmen without a cream band 4

4. (3) Vertex with lateral margins greatly elevated; its width only twice as great

as length; hind tibia with one lateral spine juscomaculosum n. sp., p. 485

Vertex over twice as wide as long ; lateral margins moderately elevated ; hind

tibiae with two lateral spines vunctiferum Walker, p. 487

5. (2) Tegmen slender, almost twice as long as broad-; strongly marked in con-

trasting light and dark patches; back concave, but no pyramidiform eleva-
tions of the tegmina wium Ball, p. 489

Tegmen short and broad; more uniformly colored, varying from cream with
few black spots to dark brown or gray with many dark spots; some form
of pyramidiform elevations on clavus 6

6. (5) Slightly larger insects measuring 3.5 to 4 mm. in length; frons with a single

median longitudinal disk narrower than either lateral disk; pronotum

longer and with margins against eye elevated bufo Van Duzee, p. 491

Slightly smaller insects, measuring 3 to 4 mm. in length; frons with a single
median disk wider than a lateral disk ; pronotuVn shorter and lateral
margins not elevated against eye 7

7. (6) Vertex with anterior margin concave through middle, making its length at

this point only one half the length of the lateral margin; angle of vertex
and frons above eye sharp; and from above this angle in position anterior
to front line of eye; claval elevations prominent sepulchralis Ball, p. 492

Vertex with anterior margins broadly, sinuately convex, length at median line
not much shorter than at lateral margin; angle of vertex and frons round-
ing; and from above this angle in position posterior to front line of eye;
claval elevations, especially at apex, scarcely visible or lacking 8

8. (7) Pale creamy in color; elevations on clavus .scarcely noticeable; size smaller,

3 mm comutum var. comutum Meliehar, p. 493

Much darker, nearer like sepulrhralis in color; claval elevations slightly longer

and more distinct; size 3 to 3.5 nun cornutinn var. iitahnum Ball, p. 495

Hysteroptennn monnn Van Duzee

Van Duzee, E. P. Expedition Cal. Acad, of Sri. to Gulf of California. Proc. Cal. Acad.
of Science 12; p. 191, 1923.

Comparative notes. The author has not seen this species. Ap-
parently it is a rare species for this country. Van Duzee described
it from one female taken at the bay at the southern end of Tiburon
Island, in the Gulf of California, and from one male taken on the
Rinco mountains, in Arizona.

Van Duzee himself distinguishes it by saying that it is a large
blackish species with much the aspect of Picumna ovatipennis
Walker, but that it is distinct structurally. He gives the size as
being from 6 to 7 mm. in length. In his description he also points
out that it resembles Hysteropterum fowleri Fowler, a Mexican
species, but is darker than that species with a more convex clypeus.

Doering: The Subfamily Issinae 483

Mr. Paul Oman, in a note to the author, states that H. morum
is close to Thionia naso Fowler, but has reticulate elytra and a
shorter vertex than that species has.

Location of types. Female is No. 1805, ]\Ius. Cal. Acad. Sci. Male
is in collection of E. P. Van Duzee.

Hysteropterum auroreum (Uhler), 1876

(Plates LVII, LVIII, LIX)

Uhler, P. R. List of the Hemip. of the Region West of the Mississippi River, Including
Those Collected During the Hayden Exploration of 1873. Bull. V. S. Geol. Geog. Surv. I,
p. 352, 1876.

Comparative notes. Superficially this species closely resembles
H. punctiferimi, both in color and size. The ground color in this
species is a brighter yellow and the markings of dark on the tegmina,
especially on the clavus, are more in the nature of blotches than dots
as in punctiferimi. The size of both species is 5 mm. Structurally
auroreum differs from punctiferum by the following characteristics:
in the former the frons, as viewed from above, shows beyond the
vertex for almost half the length of the vertex, while in the latter it
is scarcely visible; in auroreum only the median carina of the frons
is conspicuous and the rest of the disk is smooth and even slightly
convex, while in punctiferum three carinae are distinctly visible
and the disks between are concave; in auroreum the basal margin
of the frons is almost straight and the apical one deeply emarginate
to receive the large clypeus, but in punctiferum the basal margin is
convex, and the apical margin much less indented.

Male genitalia. The genitalia of this species very easily dis-
tinguish it from any allied species. The anal flap (10th abdominal
segment) usually reaches only to the tips of the styli, so that from
a ventral view it is not visible beyond them. From a flattened
dorsal view the 10th segment appears much narrower through the
basal, flattened tubular part; thence the expanded ventral margin
broadens fanlike into a fanlike flap which extends caudad a distance
which equals the length of the basal tube, after which its margins
abruptly constrict to a rounded, narrower apex. The eleventh seg-
ment shows only as a narrow rim beyond the dorsal margin of the
anal tube and bears an abbreviated stylus.

The harpagones (genital styli) are visible externally as sharply
pointed triangular plates. From this external ventral view each
harpago is twice as long as its width at base. From a flattened

484 The University Science Bulletin

lateral view each harpago is roughly pear-shaped. Its narrowed
anterior end is the point at which it is attached to the body.
Through the apical region it is broadened to over twice its width at
base. The posterior dorsal angle is prolonged cephalad into a
sharply pointed, slightly recurved hook. At the base of the ex-
panded dorsal corner is a short, broad, triangular, external hook
whose apex bends caudad.

The theca in this species closely ensheathes the aedeagus as a
tubular sheath for about two thirds its length. At this point it
splits on each side; the dorsal half then extends caudad to the full
length of the aedeagus and becomes bulged out at the apical end
into a bulbous flap, one on each side, lying above the aedeagus; the
ventral part of the split theca extends below the aedeagus as a
bluntly tapering single flap which ends in a point some distance
anterior to the apex of the aedeagus. The aedeagus shows only be-
tween the split ends of the theca as a thickly tubular structure
which bears at its extreme apex on each side a well-sclerotized
triangular hook, only part of which is visible externally, and tw^o
conspicuous, long, well-sclerotized processes which emerge, one on
each side, from the notches in the theca and extend anteriorly to the
base of the aedeagal structure.

Female genitalia. In this species, as in others in the genus, the
external female genitalia are of some taxonomic importance. The
anal flap (10th abdominal segment) is large. It extends well be-
yond the tips of the ovipositor valves and in many cases folds
ventrad around the valves, completely liiding their tips. From a
flattened dorsal view it appears as a broad wedge-shaped flap with
its lateral margins strongly, outwardly curved and with the tubular
basal region about half as wide as the expanded apical flap. The
eleventh segment is represented as a minute ringlike structure, only
barely visible beyond the dorsal margins of the tube. It bears a
small, pointed stylus.

The eighth abdominal segment is hidden under the seventh. The
ninth segment is conspicuous. The seventh segment has its posterior
margins concavely rounding and its anterior margin angulately pro-
duced cephalad.

Notes on distribution. In the original description Uhler states
that this species inhabits Texas. The specimens available for study
were all from Texas.

Location oj ttjpcs. In the United States National Museum at
Washington, D. C.

Doering: The Subfamily Issinae 485

Hysteroptenini fuscomaculosum n. sp.

(Plates LVII, LVIII, LIX)
ORIGINAL DESCRIPTION

Size. Length of body from apex of head to tip of tegmen, 2.8 mm.
to 4.3 mm., averaging around 3 mm. Length of tegmen, 2.4 mm. to
3.6 mm. Width of tegmen, 1.4 mm. to 2 mm.

Color. Same general color as punctiferum Walker. Body color
amber yellow. Vertex with lateral and anterior margins etched in
dark brown, a faint median dark-brown line, a dark spot in each
lateral half against the anterior border. Eyes mottled brown.
Disks of frons between carinae uniformly dark brown or somewhat
mottled, lateral disks lighter, mottled with dark spots; carinae and
part of lateral margins light. Clypeus brownish through middle,
crossed with darker oblique bands, median carina and anterolateral
corners light. Pronotum yellow, median light stripe, rest dotted
with brown spots. Mesonotum with borders and apex light, rest
fuscous-brown. Tegmina uniformly amber-yellow, conspicuously
dotted with small round brown spots in the cells, the veins light.
Prothorax and mesothorax tannish to brownish with margins always
light. Metathorax yellow or green. Legs amber or light yellow,
coxae and tibiae with longitudinal brown stripes or markings, femora
crossed by two to three transverse brown bands. Abdominal seg-
ments green or yellow, with usually a median brown spot against
anterior margin and two transverse rows of uniformly round dots
across disks of each segment, or else a cluster of round spots through
middle.

Structural characteristics. Shape of punctiferum Walker, but with
head and thorax proportionally narrower in comparison with great-
est width across the combined tegmina. Vertex somewhat wedge-
shaped with greatest width across its anterior margin, its width at
this point exactly twice greater than the length at lateral margins,
its posterior margin elevated. Frons narrowed at base, its basal
margin moderately convex, a distinct median carina present and two
slightly arching lateral carinae, each median disk twice as wide at
basal end as either lateral one, but about equal in size with each
lateral one at extreme apical end, apical margin deeply emarginate
to receive clypeus. Clypeus moderately inflated with a distinct
median carina present.

Pronotum in length at middle subequal to a lateral margin of ver-
tex, greatly reduced in length behind eyes. Mesonotum slightly
longer through middle than the pronotum, a transverse crease par-

486 The University Science Bulletin

allel with and close to anterior margin, a diverging oblique carina
on each lateral disk. Tegmina not quite twice as long as its greatest
width, which is at a point halfway of complete length of the insect,
slightly more inflated at this point than in acicidatum; anal margin
straight, posterior margin obliquely sloping from apex of clavus,
costal cell and part of Sc^ inflexed against venter. Wing venation
characteristic of the genus, the longitudinal veins thick, little reticu-
lation at apex.

Male genitalia. Genital segment (10th abdominal) short and not
visible from a ventral view. It is a flattened, tubelike segment at
base and has its ventral posterior margin extended into a bluntly
tapered flap, which is about one third longer than the basal part.
The eleventh segment shows beyond the dorsal posterior margin of
the tube as a small, ringlike segment, which bears an inconspicuous,
short, fingerlike stylus.

The harpagones (genital styli) are visible externally as sharply
pointed, triangular plates which meet along their inner margins.
From a flattened lateral view each harpago is roughly pear-shaped,
very much narrowed at its anterior end, where it is attached to the
body. The posterior margin is greatly widened and has its dorsal
corner elongated dorsad into a slender anteriorly recurved hook. At
the base of this dorsal extension is a narrow, liplike, external hook,
whose free margin is directed ventrad.

The internal genitalia are bilaterally symmetrical. The theca
extends for a short distance from base as a tubular sheath for
the aedeagus, then on each side it splits longitudinally. The dorsal
part continues caudad as a flap, which through the middle third ex-
tends downward on either side as a phlange in front of the aedeagal
hook, but apically splits at the middorsal line, thus terminating on
each side in two lobes, the dorsal one of the two being a long slender
spatulate process which reaches to the apex of the aedeagus, the
ventral one a short, bluntly pointed hook. The ventral part of the
theca extends caudad as a single broad, slightly tapering plate which
very nearly extends to the apex of the aedeagus. From a lateral
view the apex of the aedeagus shows between the dorsal and ventral
lobes of the theca and a slight portion of it shows dorsad of the
theca. It shows also a well-sclerotized basal hook which emerges
from the notch in the theca and is sinuately curved cephalad.

Female genitalia. Externally the genital plates and ovipositor
valves are of taxonomic importance. Very little of the valves show
externally, being partly covered by the seventh abdominal segment,

Doering: The Subfamily Issinae 487

which is triangularly produced through the middle. The seventh
segment in this species differs considerably in shape from that of
aciculatum (see drawings). The genital segment (10th abdominal)
is elongate and extends a considerable distance beyond the apices of
the valves. The tubular basal part is short, but tiie extended ventral
flap is much longer than in other species and has its lateral and
apical margins deeply reflexed, so that through the apical region the
flap is half of the basal width. The eleventh segment shows beyond
the dorsal posterior rim of anal segment as a narrow, ringlike seg-
ment which bears a short stylus.

The eighth and ninth abdominal segments are invisible. The pos-
terior margin of the seventh is triangularly produced caudad, so that
the segment is much longer through the middle than at the sides.

Comparative notes. This species more closely resembles H. pnnc-
tijerum than any other species in the genus. For comparison of
these two species see the notes under this heading in the discussion
of H. pimctiferu.m AValker.

Location of types. Holotype male, allotype female, collected
August 2, 1930, at Likely, Fla., by R. H. Beamer. Twenty-nine
paratypes from Likely, Fla., July, 1934, and August, 1930, collected
by R. H. Beamer, and eleven paratypes from Fort Pierce, Fla., Au-
gust 30, two from Lighthouse, Fla., August 2, 1930, and one from
Estero, Fla., July 21, 1934, by the same collector. These types are
in the Francis Huntington Snow Entomological Collection at the
University of Kansas.

Hysteropterum punctiferum Walker, 1851

(Plates LVII, LVIII, LIX)

Walker, Francis. List Homop. U, p. 37G, 1851.

Uhler, P. R. List of the Hemiptera of the Region West of the Mississippi River, In-
cluding Those Collected During the Hayden Exploration of 1873. Bull. U. S. Geol. Geog.
Surv. I, p. 353, 1876. Described as H. aciculatum n. sp.

Melichar, Leopold. Monograph der Issiden (Homoptera) Alih. k. k. ZoilL-Bot. Ges. Wien,
III, pt. 4, 1906.

Comparative notes. Walker's description of this species is not
very clear. Uhler, on the other hand, gives a veiy comprehensive
and easily followed description. The writer was in some doubt as to
whether they were synonymous. Mr. China of the British Museum,
however, made an accurate comparison of some recently collected
specimens and found them to be identical with the type specimen of
Walker's punctijerum collected at St. John's Bluff, Fla.

H. panctijennn Walk, is distinguished as follows: short, robust,
minutely punctate with fuscous; its length, 5 mm. It resembles

488 The University Science Bulletin

H. auroreuyn Uhler somewhat, but differs by having the frons show-
ing only a very little beyond the vertex as viewed from above ; by
the frons being distinctly tricarinate and with its basal margin
convex and the apical margin shallowly indented to receive the
clypeus. This species is more closely allied to H. juscomaculosum
than any other. They both have the same amber color, minutely
punctate on tegmina with brown flecks. They differ structurally,
however, in the shape of the vertex, size, spines on the tibiae and in
the genitalia. In jmnctijerum the vertex is well over twice as wide
as its length through middle and its lateral margins are not elevated,
while in juscomaculosum its width is only twice its length and the
margins are elevated against the eyes. The length of punctiferum
is 4 to 5 mm.; for juscomaculosum 3 mm. The head and thorax of
punctijermn are proportionately wider than in juscomaculosum.
Punctijerum has a bispinose hind tibia and the other is unispinose.
The females of the two species show differences in the shape of the
apical abdominal segments. In juscomaculosum the seventh ab-
dominal segment is triangularly produced through middle and the
ovipositor valves are partially hidden. In -punctijerum the seventh
abdominal segment is broadly produced through middle and parts
of the ninth abdominal segment, as well as the valves, are exposed.
The anal flap (10th abdominal segment) is much narrower at the
apex, due to the deeply reflexed apical and lateral margins.

Male genitalia. The tenth abdominal segment (anal flap) is more
visible externally than in H. umim, but as in that species it is not
as conspicuous as in the female. It is a flattened tube at base,
whose ventral, apical margin is expanded into a large flattened flap.
The lateral margins of the flap are bulged out through the middle,
then are abruptly constricted to form a narrowly rounded apex.
The eleventh segment shows beyond the dorsal edge of the anal tube
as a narrow rim and bears a short, pointed stylus.

The harpagones (genital styli) are visible externally as sharply
pointed triangular plates. From this external ventral view each
harpago is about twice as long as broad. From a flattened lateral
view each one is roughly pear-shaped, narrowed at its anterior end
where it is attached to the body, very much broadened across its
posterior margin, where the dorsal posterior angle is projected
dorsad as a slenderly recurved hook. At the base of this dorsal
extension is a narrow, liplike, external flap, whose free margin is
directed ventrad.

The theca and aedeagus of this species are difficult to separate,
since the theca fits snugly around the aedeagus as a tubular sheath

Doering: The Subfamily Issinae 489

with flaplike extensions. The entire structure is bilaterally sym-
metrical. From a side view most of the aedeagus is hidden. The
theca extends for a short distance from base as a tubular sheath for
the aedeagus, then on each side it splits longitudinally. The dorsal
part continues caudad as a flap, which through the middle extends
downward on either side as a phlange in front of the aedeagal hook,
while the extreme apical third is greatly narrowed, and splits
longitudinally on the dorsal side to form two elongate, spatulate
flaps, whose dorsal margins are serrate. Ventrally the theca ex-
tends caudad as a single broad, slightly tapering plate which does
not completely reach the apex of the aedeagus. The aedeagus shows
slightly at the apex as a truncate lobe between the dorsal and ventral
flaps of the theca. It also shows a semicircular, well-sclerotized
hook which emerges from the notch in the theca near its base, bends
directly cephalad a short distance and then curves caudad and
ventrad, forming a semicircle.

Female genitalia. The anal flap of the female is more conspicuous
than that of the male. It extends beyond the tip of the ovipositor
valves to a greater degree than in H. unum. (See plate LIX, draw-
ing 16.) From a flattened dorsal view it is broader at base, where
it is a flattened tube. Beyond this tubular base the ventral margin
is expanded into a roundingly tapering spatulate flap bearing numer-
ous long, stiff hairs. The eleventh segment is inconspicuous and
ringlike, showing only to a slight extent beyond the dorsal edge of
the anal tube proper. It bears a very short, buttonlike stylus.

The seventh abdominal segment is longer than in most species.
It has a sinuate posterior margin which is convex through middle.
The lateral parts of the ninth segment are visible; the eighth is
hidden under the seventh.

Notes on distribution. Type locality is St. John's Bluff, E.
Florida. Uhler states that it inhabits Orange Spring, Fla., and
Texas. In addition to many Florida specimens the writer had
available for study specimens from Okefenokee Swamp, Georgia.

Location of type. British Museum of Natural History.

Hysteropterum unum Ball

(Plates LVII, LVIII, LIX)

Ball, E. D. New Genera and Species of Issidae (Fulgoridae). Proc. Biol. Soc. Wash.
XXin, p. 43, 1910.

Comparative notes. This species is a strikingly marked species.
The vertex is light with a pair of brown V-shaped marks in the
posterior angles. The front is dark except for the lighter carinae,

490 The University Science Bulletin

margins and a few irregular spots. The elytra show contrasting
light and dark patches somewhat as follows: at base the clavus is
either entirely dark or bears an irregular brown capital L, posterior
of which the entire apex is creamy white, forming, when combined
with other tegmen, a conspicuous diamond-shaped marking in the
middle of the combined tegmina; an oblique, arcuated dark-brown
band takes up most of the corium; many of the lighter cells with
dark specks in them.

Structurally the species is characterized by the vertex, which is
not produced beyond eyes, but is two and one-half times wider than
long; the frons is tricarinate and has its basic margin deeply con-
vex; the clypeus is not deeply dovetailed into the frons and bears
a strong median carina; back concave through middle; elytra longer
than in most species, being not quite twice as long as broad.

Male genitalia. The anal flap (10th abdominal segment) is much
less conspicuous than that of the female. At base the tube is some-
what constricted, then enlarges through middle, from whence it
again narrows to a truncately rounded apex. The eleventh abdomi-
nal segment is ringlike, showing only slightly beyond the dorsal
posterior edge of the tenth segment, and bears a short, pointed apical
stylus.

The harpagones (genital styli) are visible externally as broad,
triangular lobes that meet at middle in a straight line. From this
ventral view each harpago is about one third longer than its width
at base. From a flattened lateral view (see plate LIX, drawing
13) each harpago is a somewhat pear-shaped plate, which is at-
tached at its anterior end only where it is narrowed considerably.
It is very much broadened across its apex and has its dorsal, apical
corner prolonged cephalad into a slightly cephalad-curving spatu-
late lobe, at the base of which is an external, raised, liplike flap.

The aedeagal structure is unique for the genus. It is bilaterally
symmetrical. The theca forms a close-fitting tubular sheath extend-
ing practically the entire length of the aedeagus. On each side the
apical half of the theca bears a longitudinal slit, from which arise
the aedeagal hooks. The anterior aedeagal hook arises from the
base of the slit and extends anteriorly as a forked process, the
dorsal arm of which is shorter than the ventral one. The posterior
hook of the aedeagus arises from the extreme apex of the aedeagus
as a broad spatulate flap which extends dorsad and somewhat
anteriorly. (See plate LVIII, drawing 13.)

Female genitalia. The anal flap of the female is large and very
conspicuous. It is held vertical to the body so that it very nearly

Doering: The Subfamily Issinae 491

meets the ovipositor valves on the ventral side of the body. From
a flattened dorsal view it shows its lateral margins evenly rounding
to a broad, rounded apex. The eleventh abdominal segment is not
visible as a rule externally. The stylus of this last segment is long
and slender.

The seventh abdominal segment is the last visible abdominal
segment. It is characterized by having a sinuate posterior margin
which is slightly concave through middle.

Notes on distribution. This species was described by Dr. E. D.
Ball from a single male taken near Pueblo, Colo. Specimens were
available for study from Socorro county, Carlsbad, Alamagordo, and
Rodeo, New Mexico, and from Marfa and Brewster county, Texas.

Location of types. Collection of Dr. E. D. Ball, Tucson, Ariz.

Hysteropterum bufo Van Duzee, 1923

(Plates LVII, LVIII)

Van Duzee, E. P. Expedition to the Gulf of California. Proc. Cal. Acad, of Science
12: p. 92, 1923.

Ball, E. D. Some New Issidae, With Notes on Others. Bull. Brook. Ent. Soc. XXX
(2): 37, 1935.

Comparative notes. This species measures 3.5 to 4 mm. in length.
The following color description is given in the original description:
"Color, most variable, grayish testaceous and fuscous, sometimes
nearly black or varied with green on head and pronotum, usually
with a paler area at apex of front ; elytra pale or brown with fuscous
veins and dots in the areoles." Structurally, according to Doctor
Van Duzee, it is allied to cornutum, but is distinguished by its nar-
rower front and longer pronotum. The writer finds that it is
readily separated from both cornutum and sepulchralis, which it
closely resembles, by the following characteristics: it is larger than
either of these species, being usually around 4 mm. long; the angle
formed by the union of the vertex and frons is more acute and
prominent; the pronotum is longer and has its margins against the
eye elevated; the elevations on the clavus are more conspicuous;
and the longitudinal carinae on the frons are closer to the median
one so that on each side the median disk is distinctly narrower than
the lateral one.

Genitalia. No male specimens were available for study.

The anal segment of the female very closely resembles that of
H. sepulchralis. The seventh abdominal segment is of the same
shape as cornutum and sepulchralis.

Notes on distribution. The type specimens were collected in the

492 The University Science Bulletin

Gulf of Mexico region on Lycium. Dr. E. D. Ball (1935) gives the
first North American record for this species. He took a single
female at High Tanks (Tinajas Altas) in Arizona, near the Mexi-
can border. Doctor Ball sent the writer a female to study from
Kino Bay, Mexico.

Location of type. Holotype male and allotype female in the
Museum California Academy of Science.

Hysteropterum sepulchralis Ball, 1935

(Plates LVII, LVIII, LIX)

Ball, E. D. Some New Issidae With Notes on Others. BulL Brook. Eiit. Soc. XXX
(2) p. 37, 1935.

Comparative notes. This species resembles two species rather
closely, namely, H. bufo Van Duzee and H. cornutimi var utahnum
Ball. From bufo Doctor Ball points out that it can be distinguished
as follows: "Smaller, darker, with less elevated lateral carinae of
vertex and the median tablet of the front broader. Dark brown or
gray. Length, 3-4 mm." The pronotum of bufo is also much
longer.

To distinguish this species from the species cornutum see the dis-
cussion under this heading in the description of that species.

Male genitalia. The anal segment (10th abdominal segment) is
similar to that of cornutum, although in some specimens it is a trifle
shorter and does not reach to the tip of the genital styli. From a
flattened dorsal view the anal segment is an elongate lobe, the basal
half of which comprises a flat tube, while the apical half is an ex-
tension from the \'entral margin of this tube in the form of a round-
ingly pointed flap. The eleventh segment is an inconspicuous ring-
like segment which shows only slightly beyond the dorsal posterior
margin of the tube and bears a short, slender fingerlike stylus.

The harpagones (genital styli of authors) are visible externally
from the ventral view as two triangular plates which meet along the
middle. From a flattened lateral view each harpago is roughly
pear-shaped. The narrowed anterior end is the point at which it
is attached to the body. Across its apex it is broadened to many
times its width at base. Its dorsal posterior angle is prolonged
cephalad into a curved, handlelike process. At the base of this ex-
panded dorsal corner is located a broad, bluntly rounded external
hook whose apex is directed caudad.

The internal genital structure is of the same general pattern as
that of cornutum, but differs in certain details which can best be
seen by comparing the figures on plate LVIII. The paired dorsal

Doering: The Subfamily Issinae 493

apical lobes of the theca are longer than the aedeagus in sepulchralis
and their apices are directed caudad instead of dorsad as in cor-
nutum. The single ventral apical plate of the theca extends almost
to the apex of the aedeagus in this species and the theca does not
split at the sides until it reaches a point midway of its length. The
well-sclerotized lateral hooks of the aedeagus in this species emerge
from the notch in the theca and are somewhat S-shaped as in
cornutum. However, the shape of the S is different in the two
forms; in sepulchralis the base of the hook is slender, not deeply
bent, thus making the apical loop of the figure the largest, while in
cornutum the base of the S is the largest part of the figure, due to
its broad curvature at this point.

Female genitalia. The anal segment (10th abdominal segment)
is a long slender flaplike structure which lies in a vertical plane to
the body, so that its apex touches the venter of the eighth abdominal
segment and thus almost completely hides the valves of the oviposi-
tor. This segment is longer in this species than in cornutum. It is
slightly broader through the tubular basal end than across the flap-
like apical region. The apex of the flap is roundingly pointed and
bears long stout hairs.

The seventh segment has a straight posterior margin as in cor-
nutum.

Notes on distribution. The type specimens were collected by Dr.
E. D. Ball at Bisbee, Ariz. Paratypes were also taken at Naco and
Tombstone, Ariz. All of the type material, according to Doctor
Ball, were collected on Flourensia cernua (tar-bush).

The writer had available for study a long series from the Mustang
mountains in Arizona, one specimen from the Huachuca mountains,
Arizona, and a few from each of the following places in New
Mexico: Organ, Rodeo, and Santa Fe.

Hysteropterum cornutum var. cornutum Melichar, 1906

(Plates LVII, LVIII, LIX)

Melichar, Leopold. Monograph der Issiden (Homoptera). Abh. k. k. Zool.-Bot. Ges.
Wien, in, pt. 4, 1906.

Comparative notes. Typical specimens of H. cornutum are rather
easily recognized because of its small size, since its length from apex
of head to the tip of tegmina is 3 mm.; and by its pale, uniform
creamy-yellow color of body and wings with very few dark spots
or markings. Certain specimens, however, may have brown dots
in the cells of the tegmina and when they do show this color varia-
tion then they approach rather closely the var. utahnum Ball.

494 The University Science Bulletin

Structurally comutum is allied to H. sepulchralis Ball and H. bufo
Van Duzee. It is separated from biifo by its smaller size, its pale
color, by having the single median longitudinal disk of the frons
wider or as wide as either lateral disk, the lateral margins of pro-
notum not elevated against eye and the body not concave across
back as in that species.

It is distinguished less easily from sepulchralis. The following
structural differences can be noted: in sepulchralis the anterior
margin of the vertex is concave through the middle, so that the
length of the vertex at middle is one half the length of the lateral
margin, while in comutum this margin sinuately curves cephalad, so
that the length of the vertex at middle is not much less than
at the sides; the angle above the eye formed by the union of the
vertex and frons is acute and prominent in sepulchralis, but is
rounding and inconspicuous in comutum; and lastly the claval
elevations are more prominent in sepulchralis than in comutum,
where they are usually scarcely noticeable or even lacking entirely.

Male genitalia. The tenth abdominal segment (anal flap) reaches
to the tips of the genital styli and in some cases slightly beyond, so
that the tip of it is visible beyond the apices of the styli from a
ventral view. From the flattened dorsal view the anal segment is
an elongate lobe, the basal half of which comprises a flat tube, while
the apical half is an extension from the ventral margin of this tube
in the form of a roundingly pointed flap. A small, ringlike segment,
extending beyond the dorsal margin of the tube, represents the
eleventh segment. This segment bears a slender, pointed stylus.

The harpagones (genital styli of authors) are visible externally
from the ventral view as two triangular plates which meet along
the middle. From this view each harpago appears to be about one-
third longer than its width at base. From a flattened lateral view
each harpago is roughly pear-shaped. Its narrowed anterior end
is the point at which it is attached to the body. Through the apical
region it is broadened to over twice its width at base. The posterior
dorsal angle is prolonged cephalad into a blunt spatulate lobe. At
the base of this expanded dorsal corner is a broad, liplike external
hook.

The internal genital structure is a bilaterally symmetrical tubular
organ composed of a sclerotized aedeagus surrounded almost com-
pletely by the theca. The theca surrounds the aedeagus at base as
a tubular sheath for about one third its length, then on each side
it splits longitudinally. The dorsal apical two thirds is again

Doering: The Subfamily Issinae 495

divided so that on each side it extends above the aedeagus as a long
slender lobe that reaches to the tip of the aedeagus and terminates
in a blunt recurved hook. The ventral part of the split theca ex-
tends along the lateral and ventral sides of the aedeagus as a bluntly
tapering single flap, which does not quite reach to the apex of the
aedeagus. From the notch at the point where the theca splits, a
well-sclerotized aedeagal hook emerges, and extends anteriorly and
ventralad in the fonn of a figure S and finally tapers to a finely
pointed apex.

Female genitalia. In this species the anal segment is propor-
tionally very long and the valves of the ovipositor appear shorter, so
that the result is that the latter are usually almost hidden from a
ventral view by the overlapping anal flap (10th abdominal seg-
ment). From a flattened dorsal view the flap is parallel-sided and
terminates in an evenly rounded apex. The eleventh segment is
visible only as a small ringlike segment, only slightly showing
beyond the dorsal, posterior margin of the tenth segment. It bears
an inconspicuous short pointed stylus.

The eighth and ninth segments are invisible. The seventh seg-
ment has an almost straight posterior border.

Notes on distribution. The type specimens were collected at Los
Angeles, Cal., in July, by Coquilett. Specimens were at hand for
study from the following localities: Lamar, Colo.; Carson City,
Nev. ; Emery county, Utah; St. George, Utah, all collected by David
Fox on Artemisia tridentata; Hollister, Idaho, by David Fox on
Artemisia sp.; and Cajon Pass, California.

Van Duzee (1908) stated that this species was taken by him in
abundance in Colorado and Utah.

Hysteropterum cornutvm var. utahnum Ball, 1935

Ball, E. D. Some New Issidae with Notes on Others. Bull. Brook. Ent. Soc. XXX:
38, 1935.

Comparative notes. This variety is much darker than the typical
form; in this respect it resembles more nearly H. bufo or H. sepul-
chralis. Doctor Ball describes the color as follows: "Color gray
with dark dots and spots throughout, especially marked on the
nervures and sometimes forming a double row of dots across the
anteapical cells as in sepulchralis."

Notes on distribution and host plants. Doctor Ball collected the
type series near the Grand Canyon in August, on Artemisia cana
Pursh (black sage). The writer had available for study a large

496 The University Science Bulletin

series of several hundred specimens from the Grand Canyon, Santa
Rita mountains, Flagstaff, Hereford. Coconimo county, Yavapai
county, and Oak Creek canyon, in Arizona; from Las Cruces, Blue
Springs, and Datil, in New Mexico; from Dolores and Durango, in
Colorado; and from Emery and Cove Fort, Utah.

The Genus Dictyonia Uhler, 1889

Uhler, P. R. New Genera and Species of American Homoptera. Trans, of Md. Acad,
of Science, p. 40, 1880.

Comparative notes. This genus resembles Dictyssa closely. Yet
it differs from that genus by having the tegmina practically as wide
as long, exceptionally few and large cells in the wing, the head
deeply sunkeii so that the eyes almost touch the base of the tegmina,
and the terga of the abdominal segments so compressed that a
conspicuous, elevated and somewhat swollen ridge is formed down
the middle of the abdomen.

Like the genus Dictyssa it has bispinose hind tibiae and small or
rudimentary hind wings.

The genus also superficially resembles the genus Neaethus Stal,
but it differs from the latter in the reduced number of the cells
and their exaggerated proportions on the central area of the tegmina.

Dictyonia obscnra Uhler, 1889

(Plates LXVI, LXVII)

Uhler, P. R. New Genera and Species of American Homoptera. Trans, of the Mary-
land Acad, of Science, p. 40. 1889.

Comparative notes. This genus is monotypic and therefore the
distinctive characteristics listed above for the genus are also used
in identifying the single species obscura. Specifically Dictyonia
obscura resembles Dictyssa aereolata Melichar and Dictyssa qua-
dravitrea n. sp. more than other species in the subfamily. Fre-
quently in collections it has been determined as the former. In
addition to the generic differences it is easily distinguished from
D. aereolata by the angular shape and large size of the cells of the
tegmina, the widely expanded costal margin and the presence of
the broad, cream-colored band down the middle of the frons. It
is distinguished externally from D. quadravitrea by the extreme
broadness of its tegmina, the wide expanded costal margin and
again by the cream-colored band on the frons.

Wing venation. In general the wing venation is of the same pat-
tern as that of the genus Dictyssa. Vein Sc^ extends along the costal
margin as a distinct vein for two thirds the length of the tegmen.

Doering: The Subfamily Issinae 497

Veins SCo and R are united for a short distance after leaving the
central trunk, then R extends as a single straight vein and Scg as an
angled vein toward the apex. Vein M divides near apex into two
short branches. Vein Cu, runs so close to claval suture that an un-
usually large hyaline cell occurs between it and vein M; Cu^
branches into two branches which are not widely separated from the
claval suture. Vein Cu. extends along the claval suture (Muir).
Veins 1st A and 2d A are united at apex to form the Y-vein of the
clavus.

Male genitalia. The tenth abdominal segment (anal flap) is an
elongate flattened tube with its greatest width at middle, where it is
about one half its length, from whence it tapers to a truncate, or
sometimes slightly emarginate, apex. The eleventh segment is much
reduced, scarcely visible in the anal tube, except for the presence of
its elongate stylus, w4iich shows as a slender projection beyond the
dorsal margin of the anal flap.

The harpagones (genital styli) are visible externally from the
ventral aspect as two adjoining, tapering plates. From a flattened,
lateral view (see plate LXVII, drawing 8) each harpago appears as
a subquadrangular plate with its dorsal posterior corner projected
dorsad as a sharply pointed extension, at the base of which exter-
nally is located a flat, recurved hooklike structure.

Like many other genera the aedeagus is asymmetrical. On the
left side it appears as a slender, tubular, somewhat pointed structure
which is less sclerotized at tip than at base. Attached near its base
and extending beyond the theca to a point not quite midway of
aedeagus are two parallel, equal-sized heavily sclerotized processes.
On the right side the aedeagus bears no sclerotized processes. The
tubular membranous theca surrounds the basal third of aedeagus.
On the ventral side it is truncate posteriorly, thus allowing the
aedeagal hooks to show beyond it. On the dorsal side it extends
caudad as a finely tapering projection, whose tip reaches to a point
just beyond middle of aedeagus and somewhat caudad to the tips of
the aedeagal hooks.

Female genitalia. The anal flap (10th abdominal segment) of
the female is similar to that of the male. The external valves of
the ovipositor have their greatest width through the middle, at
which point they are approximately half as long as wide.

Location of types. Uhler type is lost, according to Mr. Paul Oman,
of the National Museum, Washington, D. C.

Notes on distribution. Uhler states that this species had been col-

32—4141

498 The University Science Bulletin

lected only from central California and around San Francisco. Ap-
parently he described the species from two specimens, or at least
only a few. Doctor Ball has sent the writer specimens from Yosem-
ite and Vesalia, Cal. In the Snow Entomological collection at the
University of Kansas is a large series collected from Giant Forest,
Cal, in June, 1929, and Three Rivers, Cal., in June, 1932, by Dr.
R. H. Beamer, and one specimen from Republic, Wash., by John
Nottingham, in August, 1931. In the National Museum Collection
is a large series, collected by Paul Oman at Three Rivers, Cal., in
June, 1935, and a smaller series from Mariposa, Cal.

The Genus Dictyonissus Uhler, 1876

Uhler, P. R: List of Hemiptera of the Region West of the Mississippi River, Including
Those Collected During the Hayden Explorations of 1873. Bull. U. S. Geol. Surv. I, p.
354, 1876.

Comparative notes. The main distinguishing characteristics of
the genus are: the collarlike pronotum, which in front is acutely, tri-
angularly narrowed and carried forward almost to the front line of
the eyes and is deeply emarginated behind; the long mesonotum,
which through middle is as long as combined vertex and pronotum;
long, narrow tegmina with costal area not expanded, the veins thick
and conspicuous, making a network of large, round aereoles, so that
the course of the main longitudinal veins cannot be traced; and the
conspicuous, erect hairs on body and tegmina. Tibiae of hind legs
with 3 spines. Hind wings reduced to mere scales.

Key to the Spextibs

la. Tegmina and body sparsely covered with pitch black, long, stiff hairs; anterior

margin of vertex distinctly angulate at middle D. jiiaropilosus n. sp., p. 499

lb. Tegmina and body sparsely covered with light hairs; anterior margin of vertex

evenly rounding D. griphus L'hler, p. 498

Dictyonissiix (ir'phus Uhler, 1876

(Plates LXVI, LXVII)

Comparative notes. For comparison with D. nigropilos^is, which
it closely resembles, see the notes under this heading in the descrip-
tion of that species.

Either species of Dictyonissus might possibly be confused with
Misodema reticulata, especially if one has never seen the latter,
since both genera are described as having coarse veins making a
network in the tegmen, and the same type of vertex. The two genera
are easily separated, however, by the following characteristics:
Dictyonissus has a pale green body, pale green tegmina with liyaline

Doering: The Subfamily Issinae 490

areoles and long, erect hairs on the veins of the tegmina and the
body; Misodema has a fuscous body coloring, and the tegmina are
opaque and hairless.

Male genitalia. Anal flap (10th abdominal segment) about twice
as long as wide, parallel-margined for two thirds its length, then
tapering suddenly at the apex. The eleventh segment and its stylus
short and inconspicuous.

The harpagones as viewed externally on the ventral side are two
long, slender processes. From a flattened lateral view the dorso-
caudal angles of each is found to be prolonged dorsad into a sharply
pointed, slightly recurved apex, and at the base of this extension is
found an external ventrad-curving, sharply pointed flap.

The aedeagus in this species is partially bilaterally symmetrical.
It is tubular in shape and only partially exposed beyond the theca.
On the apical three fourths it bears a sclerotized phlange which
tapers to a narrow ridge toward the apex and which is serrate along
the dorsal margin. The theca appears as a tube covering the aedea-
gus for approximately one half the length of the latter, then on each
side it suddenly projects caudad as a narrow, elongate flap which
extends to about the base of the apical third of the aedeagus.

Female genitalia. Anal flap (10th segment) broadest through
middle, suddenly tapering to a roundingly pointed apex. The elev-
enth segment scarcely visible, its stylus moderately long. The ex-
ternal valves of the ovipositor broadest through middle third.

Notes on distribution. Uhler states that it inhabits Texas. The
type specimen was collected at Waco by Belfrage. In 1929 a large
series was collected in Kerr county, Karnes county, and Menard
county in Texas by R. H. Beamer. Several specimens were taken
in 1932 at Monterey, Nuevoleon, Mexico, by L. D. Tuthill.

Dictyonissus nigropilosus n. sp.

(Plates LXVI, LXVII)
ORIGINAL DESCRIPTION

Size. Length of body from apex of head to tip of tegmen, 3.3
mm. to 3.8 mm. Length of tegmen, 2.7 mm. to 3.2 mm.; width of
tegmen, 1.6 mm. Much smaller than griphus.

Color. Same general dight green to testaceous color as in D.
griphus, but with pitch black hairs over body and tegmina instead
of light ones. Body uniform light green or greenish-tan. Anterior
margin of vertex darkish. A bronze caste over disk of mesonotum.
Eyes brown to black. Frons without the black dots found in griphus,

500 The Univeksity Science Bulletin

but base of black hairs also sometimes black. Venter of body green.
Legs also green except for a bronze caste on tarsus, and tip of tarsal
claws black. Tegmina greenish or greenish-tan, translucent, and
very thick green veins.

Structural details. Vertex greatly protruding beyond eyes, its
total width ecjual to its total length, the anterior margin distinctly
angulately produced through middle, the posterior margin deeply
roundingly emarginate, lateral angles parallel. Frons greatly elon-
gate, narrowed at base, its lateral margins outwardly curving so
that its greatest width is at base of apical third; a distinct median
carina present for about tw^o thirds its length, the entire disk to-
gether with the carina at base considerably elevated, leaving a shal-
low groove just mesad of each lateral margin, which is also elevated.
Clypeus short, about two thirds the length of one lateral margin of
the frons. Femora of legs slightly sulcate, tibiae greatly so. Pro-
notum collarlike as for the genus, but each lateral arm more rounded
and elevated into a knob than in D. griphus. Mesonotum conspicu-
ously humped at anterior end, two distinct lateral carinae present
and a median longitudinal groove plainly visible. Costal margin of
tegmen only slightly rounding, costal cell area broad, more broadly
extended at anterior end than in griphus; longitudinal veins only
distinguishable for a short distance at base, after which they are lost
in the reticulation.

Male genitalia. Anal flap (10th abdominal segment) tubular at
base for half the length, from which point the ventral margin ex-
tends posteriorly into a bilobed, somewhat pointed flap. Beyond
the dorsal margin the ringlike eleventh segment is only slightly visi-
ble and bears a long fingerlike stylus.

Each harpago from a flattened lateral view is roughly boot-
shaped, broadest at its extreme apical end, where the dorsal corner
extends into a sharply pointed triangular projection. A flat, caudad-
bending external hook is located midway of this projection.

The aedeagus is a semisclerotized tubular process which extends
about halfway beyond the theca. On the right side it bears a nar-
row phlange which is saw-toothed on the dorsal margin. Just pos-
terior to the end of the phlange is a recurved, well sclerotized hook.
On the left side a similar phlange with a serrate dorsal edge is also
present and another well-sclerotized hook, the apex of which only,
shows beyond the edge of the theca as a sharply pointed bladelike
process. The theca envelopes the aedeagus for half its length and
has its caudal margin on both sides deeply notched through middle.

Doering: The Subfamily Issinae 501

Comparative notes. There are only two species in the genus.
They resemble each other closely in color, size and general features.
They are easily distinguished by the presence of pitch-black hairs
on body and tegmina in nigrapilosus, together with the angulate ver-
tex and the very narrow frons without any black markings. D.
griphus has golden or light hairs, the anterior margin of vertex dis-
tinctly rounding and more produced beyond eyes and a broader
frons with a row of black spots indicated in each lateral disk. D.
griphus usually is a trifle larger than nigropilosus, its measurements
being 3.5 mm. to 3.8 mm. from apex of head to tip of tegmen.

Location of types. Described from 3 males and 1 female. Holo-
type male and allotype female and two male paratypes taken at
George West, Texas, July 1, 1936, collected by R. H. Beamer.

These types are in the Snow Entomological Collection, University
of Kansas.

The Genus Neaethus Stal

Stal, Carolus. Rio Jan. Hemip. II, p. 67, 1862.

Description of the genus. This genus is similar to Dictyssa and
Dictyonia. They are small insects with hemispherical, translucent
or semitransparent tegmina and thick veins. The apical margin of
each tegmen is strongly rounded and the costal margin expanded.
Vertex is short through middle, much wider than long, not produced
greatly beyond eyes. Pronotum extremely narrow at sides, where it
is shortened to almost a point ; its anterior margin deeply emarginate
into region of the vertex; its posterior margin shallowly concave.
Mesonotum is triangular with or without a median carinae and an
arcuated groove just back of anterior margin. Frons is held ver-
tically to body, is usually more or less parallel-margined, truncate
posteriorly, and deeply emarginate anteriorly for the insertion of the
postclypeus. Clypeus is triangular, not keeled on margins. Tegmina
held vertically and lying adpressed to body, translucent or semi-
opaque, always with heavy, conspicuous veins and many cross veins
forming small angular cells. Wing venation, based on Muir, Till-
yard and Snodgrass, is similar to Dictyssa, showing the following
general characteristics: vein Sc divided, Sc^ (equal costal vein of
Metcalf ) running along costal border for about one third the length
of the tegmen, veins SCo and R united at base and their apices some-
times completely lost in reticulation, in other cases each branches
again near tip; vein M always two branched, the branches of each of
these either apically not traceable in the reticulation or frequently
branched several times, making a 4-, 5- or even 7-branched vein; vein

502 The University Science Bulletin

CUj divided into two branches, CUja and CUji,; vein Cu^ forming the
claval suture; 1st A and 2d A present. Hind wings are of three
types; as long as the tegmina, very much abbreviated, but with a
longitudinal fold indicated, or reduced to a mere scale. Two spines
are present on the hind tibiae.

HISTORY OF THE GENTS

The genus was erected by Stal in 1862. Previously he had de-
scribed vitripennis (Of. Vet. Akad. Forh. XI, p. 247) in 1854 as a
Hysteropteriim. This species is now regarded as the haplotype of
the genus Neaethus. In 1906 ]Melichar described four other species,
namely, jenestratus, grossus, nigronervosiis and maculatits. In 1921
Van Duzee added fragosus to the group. In the present paper seven
new species are being added to the list of North American species,
making a total of thirteen.

On the whole the species in this genus are not easily recognized by
external characteristics. The most closely allied species groups
perhaps are the nigronervosus-jenestratus group and the diversus-
unijormus-curvamimis group. The first one has widely different
genitalia. The latter group have somewhat similar genitalia. The
general reader may have difficulty in identifying all of these species
by external characteristics. Close study of the genitalia show dif-
ferences that are constant and it is upon these differences, since in
some instances long series of slides were made from several localities,
that the species have been erected. The home of the genus seems to
be California and the Southwest, w'here many of them have been
taken from oaks. Mr. Paul Oman, of the National Museum, and
Dr. R. H. Beamer have stated that they have taken them from
different varieties of oak trees even in the same mountain range.
Apparently this genus is one of the highly variable groups w^hich
has split up into many species corresponding to different species or
varieties of the host-plant group.

Key to Species

1. Hind wings as long as or only slightly shorter than front wings 2

Hind wings rudimentarj-, less than one fourth length of front wings 4

2. (1) Tegmina translucent and held vertically against body; pronotuni greatly

narrower than head perlucidtis n. sp., p. 503

Tegmina more or less opaque and held somewhat horizontally ; pronotum only
slightly narrower than head 3

3. (2) Larger insects, 5 to 6 mm. long; lateral margins of vertex longer than width

across eye; narrow c«staJ cell fragosus Van Duzee, p. 506

Smaller insects under 4.6 mm. ; vertex concave, short, one lateral margin

equal to or less than width across eye; broad costal cell ... .grossus Mel., p. 507

Doering: The Subfamily Issinae o03

4. (1) Tegmina translucent, with a black band extending from clavvis to costal

margin and a brown spot at apex of claviis; pronotum deeply notched

behind eye maciilatus Mel., p. 510

Tegmina without such bands or spots; pronotum not so deeply notched 5

5. (4) Translucent tegmina with light veins ^

Tegmina more opaque, with either very thick, light or darker to black, thin

7
veins '

6. (5) Lateral arm of pronotum back of eye longer, anterior margin of pronotum

notched; postclypeus length equal to one lateral margin of frons; costal
cell wide and distinctly reflexed ; vein Cu branching considerably anterior

to middle vitripennis Stal, p. 512

Lateral part of pronotum short, not knobbed or deeply notched behind eye;
postclypeus shorter; costal cell narrower and not reflexed; vein Cu
branched at about middle sinehamatiis n. sp., p. 515

7. (5) Larger insects, usually 4.4 mm. or over; vein Sci straight and long, extending

for at least one third lengtli of wing; hind wings rudimentary but folded

once longitudinally °

Smaller insects, under 4.4 mm. (occasionally females larger); vein Scj dis-
tinctly curved and short, not extending one third of wing length; hind
wings reduced to mere scales with no longitudinal folds 9

8. (7) Head produced beyond anterior margin of eye; frons narrow, one lateral

margin greater than its basal margin ; anal flap of male two-pronged

similis n. .sp., p. 517

Head scarcely produced beyond eye; frons broad, length and width about

equal ; anal flap of male rounded jacintiensus n. sp., p. 519

9. (7) Wins either all dark or at least cross veins dark or darkish; vein Cui not

uniting before apex or lost in reticulation 10

Veins light throughout ; vein Cu, uniting in some way before apex 11

10. (9) Veins of female usually all black (longitudinal veins of male and some females

lighter); vein Sci greatly arched, short, not more than about one fourth

length of tegmen; apical cells long and parallel nigronervosus Mel., p. 522

Longitudinal veins light; vein Sci short, but straight; a cluster of small

ce-lls at apex of coriuiii fenestratus Mel., p. 525

11. (9) Veins thin, costal cell wide, vein Sci distinctly inwardly arched so that

costal cells are wider tlian long; elongate cells at apex as in nigronervosus,

curiaminix n. sp., p. 528

Veins thick, costal cell narrower, with the small cells in this area either longer
than wide or square; an irregular reticulation of small cells at apex 12

12. (11) More cro.ss veins in costal cell area, numbering usually 7 or 8; clypeus

longer, its length equal to one lateral margin of frons. . . .uniformus n. sp., p. 530
Fewer cross veins in costal cell area, numbering only 4 or 5 ; clypeus shorter,

its length only equal to basal or posterior margin of frons. .diversus n. sp., p. 532

Neaethus perlucidus n. sp.

(Plates LX, LXL LXIl, LXIII)
ORIGINAL DESCRIPTION

Size. Length of body to tip of tegmen, 4.4 mm. to 4.6 mm.
Length of tegmen, 3.6 mm.; width of tegmen, 2.6 mm. This is a
medium-sized species for this genus.

Color. Uniform straw-yellow, with few darker markings. Body
uniformly a darker or straw-yellow, except margins of all segments
and sclerites light cream, also a faint cream line down middle of

504 The University Science Bulletin

vertex, pronotum and mesonotum and middle abdominal segments
from below with a greenish caste. Tegmina translucent straw-
yellow or light amber; veins same color except a few dark cross
veins on inner border of clavus and again at apex. Hind wings pale
yellow, translucent. Eyes brownish. Clavola (flagellum) of antenna
brownish-black. Legs all yellow except for black tips of the spines.

Structural details. Vertex subrectangular, slightly concave, much
shorter through middle than at sides, its width at the anterior
margin twice the length of one lateral margin, the lateral margins
outwardly convex. Greatest width of eyes half the width of vertex.
Frons depressed lengthwise through middle, lateral margins elevated,
and slightly outwardly convex so that greatest width of frons is
through its anterior fourth, median carina present but disappearing
before reaching clypeus. Postclypeus with posterior margin angu-
larly produced into frons for a distance equal to about one fifth the
length of the latter.

Pronotum much narrower than head, its anterior margin round-
ingly produced for greater distance into vertex than in I'itripennis,
its posterior margin shallowly emarginate; its extreme lateral part
behind eye greatly abbreviated to about one sixth its length through
middle and not deeply notched as in maculatus. Mesonotum with
only a very faint median carina present, a deep transverse groove
following anterior margin, but stopping at either side before reach-
ing lateral margin, where each lateral corner is bullate. Tegmina
approximately one third longer than wide, their greatest width ap-
proximately through middle, hemispherical in outline, due to both
the apical and costal borders being evenly rounding; longitudinal
and cross veins heavy, and forming large, irregular, angulate cells;
costal cell area very broadly expanded; vein Sc^ running as a stiff,
straight vein from base to costal margin, making the cells in middle
of the expanded border much larger than either the anterior or
posterior ones; vein M only four-branched; vein Cu branched at
about middle and veins Cu^a and Cu^b united before apex again
and the combined vein shortly uniting with vein M3 ^ ^. Hind
wings as long as tegmina.

Male genitalia. Anal flap (10th abdominal segment) in flattened
view approximately twice as long as wide with its lateral margins
subparallel and its posterior margin roundingly emarginate. Elev-
enth segment ringlike, only slightly showing beyond rim of tenth
segment and bearing the usual fingerlike stylus.

The harpago (genital stylus) from a flattened lateral view ap-

Doering: The Subfamily Issinae 505

pearing gubqiiadrangular, broadest at its middle, due to slight bulg-
ing at this point of the ventral margin, the dorsal posterior corner
extended dorsad into a sharply pointed process, at base of which is
a ventrad-curved small external hook.

The aedeagus as viewed from the left side is a partially sclerotized
tube bearing three sclerotized pointed flaps located as follows: a
short curved one attached approximately at apex on anterior fourth
and which is gently curved dorsad, a long, flat, sharply pointed one
attached somewhat near base of aedeagus so that only its distal half
shows beyond the theca, and the third attached near the middle and
extending to near the apical curve of the aedeagus. The apical
ninth of the aedeagus is recurved and is bulbous at base, then nar-
rows to a sharply pointed apex. On the right side the aedeagus
shows only one sclerotized hook or flap whose apex reaches ap-
proximately to base of apical third. The semisclerotized theca
covers the aedeagus at base. On the left side it appears as a bluntly
pointed sheathlike flap. On the right side it extends about one
half the length of the aedeagus. Posteriorly it is truncate on the
dorsal side and on the ventral side extends posteriorly as an elongate,
slightly dorsad-curving hook.

Comparative notes. This species has probably always been
identified as A^. vitripennis Stal, because of its close resemblance
in size, coloring and translucent tegmina. For the same reason it
would be confused with sinehamatus n. sp. It is readily separated
from these two species by the presence of long hind wings. For
further details of comparison, see the notes under this heading in
the descriptions of these species.

The presence of long hind wings places it near fragosiis Van
Duzee and grossus Melichar. It is easily distinguished from these
two by the transparency of the tegmina and the lack of maculation
or dark veins which are found on them. Other characters which
distinguish these three species are as follows: perlucidus has a much
narrower thorax than either of the other two; perlucidus and
fragosus have the pronotum deeply produced anteriorly into the
vertex, while grossus is only shallowly produced; in perlucidus the
expanded costal border is very broad, and vein Sc^ is very straight,
thus making the cells in the middle of the costal area larger than at
either end; in grossus Sc^ curves outwardly, following the costal
border, and thus the transverse cells are practically the same width ;
in fragosus the costal area is expanded only half as much as in
grossus and perlucidus; grossus is conspicuously gibbous between

506 The University Science Bulletin

Sci and R just in front of middle, which condition is only faintly
indicated in fragosus, entirely lacking in perlucidus; finally the
position in which the tegmina are held against the body varies in
the three, in fragosus being held more horizontal, in perlucidus
almost vertical and in grossus halfway between the other two.

Notes on distribution. Collected at Three Rivers, Cal., on July
8, 1932.

Location of types. Holotype male, collected at Three Rivers,
Cal., July 8, 1932, and three paratype males same place, by Dr.
R. H. Beamer. These types are in the Snow Entomological Collec-
tion at the University of Kansas.

Neaethus fragosus Van Duzee, 1921

(Plates LX, LXI, LXII, LXIII)

Van Duzee, E. P. Characters of Some New Species of North American Hemipterous
Insects with one New Genus. Proc. Cal. Acad, of Sci. XI (10) pp. 111-134, 1921.

Comparative notes. One of the largest species in the genus,
measuring 4.5 to 6 mm. from apex of head to tip of tegmen. Readily
distinguished by the semiopaque, long and narrow tegmina with a
narrow, expanded costal border; vein M 5-branched; veins Cu^a
and Cujb united before apex; and the presence of hind wings which
are nearly as long as the tegmina.

Van Duzee gives a description of the coloration as follows:
"Yellowish or greenish testaceous, in male usually becoming brown-
ish on vertex and face, with a paler area on apex of front; veins of
elytra sometimes distinctly infuscated, especially in the male."

This species closely resembles N. grossus Melichar. It also is
placed near N. perlucidus n. sp., because of the long hind wings.
For detailed comparison of these three species see the notes under
this heading in the description of N. perlucidus.

Male genitalia. Anal flap (10th abdominal segment) compara-
tively broad, its lateral margins outwardly curving to about their
middle, then converging to a narrowed truncate apex, the flaplike
extension from the basal margin not ciuite half the length of the
entire segment. Eleventh abdominal segment ringlike with a short
fingerlike stylus.

Harpago as viewed from a flattened lateral view roughly sub-
cjuadrangular, its ventral margin rounding outwardly, its dorsal
margin shallowly convex through middle, but at apical third ex-
tending dorsad a short distance, then making an obtuse angle and
running posteriorly. The extreme dorsoposterior corner extended
posteriorly as a short, blunt projection, of which both the ventral

Doering: The Subfamily Issinae 507

and posterior corners are finely pointed and somewhat recurved.
At the base of this posterior projection is a slender, recurved ex-
ternal hook. The aedeagus is somewhat more complex in this
species than in others of the genus. Its basal half is covered by the
theca. The apical portion that is visible beyond the theca is curved
into a semicircle, is somewhat membranous so as to appear slightly
wrinkled on the dorsal side and tapers to a broadly rounded apex.
From a left view^ two well-sclerotized processes are visible; one a
broad, flat projection attached at middle of aedeagus and extend-
ing caudad to about base of apical sixth, where it ends in a finely
pointed apex; the other a stout, hornlike process attached nearer
to base of aedeagus and only partially visible beyond the posterior
apex of the theca. This hook is broad at base, but gently tapers to
a pointed apex. It runs parallel with the aedeagus for a short dis-
tance, then, just after its emergence from the theca, it abruptly bends
dorsad toward the aedeagus and partially overlaps the rim of the
latter on the right side. This ventral hook is visible from either
side of the aedeagus. On the right side there is present a similar
dorsal flat projection as is seen on the left. The theca is practically
the same on either side. It appears basally over the aedeagus as a
membranous tube. At its dorsoposterior corner it extends posteriorly
as a rounded elongate flap which ends in a small short dorsal spine.

Notes on distribution. The type locality is the summit of Mt.
Wilson, Pasadena, Cal. A series of twenty-five specimens have
been collected by Dr. R. H. Beamer from the following places in
California: Anza, Idyllwild, Three Rivers, and San Jacinto moun-
tains, in July and August.

Location of types. Museum of the California Academy of Science.

Neaethiis grossus Melichar, 1906

(Plates LX, LXI, LXII, LXV)

Melichar, Leopold. Monograph der Issiden (Homoptera). Abh. k. k. Zool.-Bot. Ges.
Wien, III, pt. 4, 1906.

Size. Length of body to tip of tegmen, 4 mm. to 4.6 mm. Length
of tegmen, 3.37 mm. to 3.8 mm.; width of tegmen, 2.08 mm. This
is one of the median-sized species in the genus.

Color. General color yellow, mottled with blackish-brown. Eyes
yellow, spotted with red or all reddish-brown. Vertex dark tan with
a cream-colored carina down middle, lateral margins also cream,
enlarging to a spot at posterior corner. Pronotum tan with a cream-
colored median carina and a cream spot at each corner behind eyes.
Mesonotum brown except for two yellow spots on each side, one of

508 The University Science Bulletin

which lies anteriorly against the anterior margin and the other in
line with it but on the posterior border, apex also cream. Frons
reddish-brown, thin outer margins dark brown with a narrow white
band bordering them on the inside and a large spot in center which
is cream. Postclypeus reddish-brown, anteclypeus yellow, a dark
longitudinal abbreviated stripe on each side running across both
sclerites. Venter of body light yellow through middle, becoming
darker at the sides and fifth and sixth abdominal segments washed
in brown through middle and at extreme sides. Ventral ovipositor
valves red-brown with their inner margins forming a dark-brown
line. Legs brownish through middle, lighter at apices of segments,
claws on tarsus black-tipped. Tegmina ground color semiopaque
creamish-yellow, longitudinal veins yellow, cross veins varying from
light brown to dark brown, usually the dark brown ones forming an
irregularly spotted band, starting on clavus in line with apex of
mesonotum and extending vertically across corium to costal border
or sometimes forming more of an elongate vertical spot rather than
a band, many dark veinlets again indicated in irregular patches in
posterior half.

Structural details. Vertex subrectangular, broad, being at least
four times wider across anterior margin than at middle, lateral
margins diverging posteriorly, each margin twice as long as length
through middle, anterior margin almost straight, only slightly pro-
duced anteriorly, not much greater than anterior margin of eye.
Frons depressed through middle, its lateral margins distinctly ele-
vated and diverging anteriorly so that greatest width is through its
anterior fifth, a median carina distinctly indicated for two thirds
its length. Pronotum scarcely narrower than head, roundingly and
comparatively shallowly produced anteriorly and narrower at this
point than in maculatus. Mesonotum one third wider than long,
each lateral fourth slightly bullate, a deep crescent-shaped groove
following the anterior margin, but stopping just mesad of each
lateral bulla; a very faint median carina present. Tegmina ap-
proximately one third longer than wide, their greatest width in line
with apical sixth of clavus, distinctly bullate between veins Sc^ and
R at basal third of wing, costal cell area widely expanded. Longi-
tudinal veins more elevated than cross veins, but of less diameter
than many of the latter, the cross veins forming an irregular net-
work of small veinlets. Vein Sc^ rather gently curving, following
outline of costal border, so that the cells in the expanded costal
region are practically the same size and the transverse veins in this

Doering: The Subfamily Issinae 509

region usually forked at end. Vein Mg^^ often 3-branched, making
a 5-branched media; all the branches of M and Cu^ posteriorly
arched and running close together, but not united before apex as in
some species. Hind wings as long as tegmina.

Male genitalia. Anal flap (10th abdominal segment) in flattened
view not quite twice as long as wide; its lateral margins subparallel
and its posterior margin roundingly produced. Eleventh segment
ringlike, only slightly visible beyond rim of 10th segment and bear-
ing tlie usual fingerlike stylus.

The harpago (genital stylus) from a flattened lateral view appear-
ing as a subquadrangular flat plate about twice as long as wide, its
ventral margin outwardly rounding, the dorsal margin slightly
convex through middle and its posterior angle prolonged cephalad
as a short pointed hooklike process, another triangular external
hook attached at base of dorsal process and with its apex directed
ventrad.

The aedeagus, as viewed from the left side, is a partially sclero-
tized tube bearing two sclerotized external processes, the more
distal one attached to middle and extending caudad as an elongate
narrow flap, which narrows to a bluntly rounding point reaching to
about the base of the distal eighth of the aedeagus; the second
process at base covered by the theca, but attached somewhat near
the apex of basal fourth of aedeagus and showing beyond the
theca as a well-sclerotized hook, w^hich is broad at base, gently
narrows to a slenderly pointed apex and reaches not quite to middle
of aedeagus. On the right side the aedeagus shows only one flap-
like extension which rises obliquely from the sides of the aedeagus,
beginning at base of apical third and whose pointed apex reaches
to base of apical sixth of aedeagus. The theca covers the base of
aedeagus as a tube, extending on the left side about one third of
the length of aedeagus, and ends posteriorly with a truncate margin.
On the right side the dorsal margin of the theca is extended caudad
as a sinuately margined flap which at its apex suddenly bends
ventrad in the form of a pointed hook.

Comparative notes. This species more closely resembles A^. fra-
gosus Melichar than any other one. It is separated easily from
the latter by having a, distinct bulla at the base of each tegmen
and by the more widely expanded costal border. For more details
of comparison see notes under this heading in the description of
N. perlucidus.

Notes on distribution. Type locality is given as Los Angeles, Cal.

510 The University Science Bulletin

In the Snow Entomological Collection are specimens from the Santa
Rita mountains, Ariz., and the following places in California: Mint
Canyon, Monrovia Canyon and Lockwood.

Location of type. Types are in the National Museum in Wash-
ington, D. C.

var. pallida Melichar

Melichar states that dark speckling or bands on the tegmina are
sometimes lacking so that they appear pale yellowish. He calls
this type variety pallida. Out of twenty-seven specimens studied
the writer found eight which would fall under this classification.

Neaethus maculatus Melichar, 1906

(Plates LX, LXI, LXII, LXV)

Melichar, Leopold. Monograph der Issiden (Homoptera). Abh. k. k. Zool.-Bot. Ges.
Wien, III, pt. 4, 1906.

Size. Length of body from apex of head to tip of tegmen is 4
mm. to 4.4 mm. Length of tegmen, 3.6 mm. to 4 mm.; greatest
width of tegmen, 2.132 to 2.532 mm.

Color. General color light tan with transparent tegmina crossed
by two oblique brownish-fuscous bands. Eyes light brown, banded
or spotted in red. Vertex usually uniform yellow, sometimes spotted
with light brown, especially females. Pronotum yellow tan with
dark, paired, round pits in middle, sometimes numerous dark dots
at sides. Frons and clypeus light yellow, a dark brown longitudinal
streak on each side just laterad of the median carina and another
irregular dark longitudinal streak more laterad to this, followed by a
border of brown dots, then a longitudinal pale band, finally bor-
dered by the very thin, dark lateral edge. Clypeus usually with
a light median carina and sides washed in brown or with numerous
oblique brown stripes on each side. Venter of body either uniform
light yellow or frequently washed in fuscous, especially across bases
of segments or at sides along the margins. Tegmina translucent
greyish-tan with conspicuous dark veins, a prominent dark-brown
oblique band before middle, beginning on base of clavus and extend-
ing slightly posteriorly across corium to a point on costal margin
where vein Sc^ ends; another interrupted band beginning at apex of
clavus extending half across corium where it is lacking for width of
one cell, then starts again somewhat more posteriorly and ends
on apical margin. The latter part of this band or spot is fre-
quently entirely lacking. Legs uniform yellow or sometimes washed
in fuscous or with longitudinal carinae fuscous; claws black tipped.

Doering: The Subfamily Issinae 511

Structural details. Vertex not much produced beyond eyes, con-
cave through middle, anterior margin straight, at this margin 41/2
times wider than length along middorsal line, lateral margins con-
verging slightly anteriorly. Pronotum much narrower than head,
anterior margin broadly rounding, deeply notched behind eye and
greatly overlapping vertex through middle; two round paired pits
in center of disk. Mesonotum one third longer than wide, three
carinae present, each lateral fourth bullate, a deep crescent-shaped
groove following the anterior margin but stopping just mesad of
each lateral bulla. Frons with basal or posterior margin straight, a
median carina present for practically the entire length, lateral mar-
gins distinctly parallel and greatly elevated. Clypeus with a median
carina present and deeply inserted into frons. Tegmina roughly
twice as wide as long, superficially wedge-shaped with greatest width
just anterior to apex of clavus; veins distinct, very few cross veins,
so that the cells of wing are angular and easily counted; vein Sci a
wide distance from costal margin, so that this species has an espe-
cially widely expanded costal area, the transverse veins in the region
numbering 8 or 9. Venation characteristic for the genus with veins
Sc,, R, M,,. and M^^^ each branching once, making a 4-branched
media, Cu^a and CUii, fused before apex and the single vein follow-
ing apical border until it runs into vein M.^^^, thus setting off a
narrow phlange. Hind wings very short or rudimentary, folded once
longitudinally. (Melichar stated that they are lacking.)

Male genitalia. Anal flap (10th abdominal segment) more elon-
gate than in many species, its length being 2i^ times its greatest
width, which is across the flap part of the segment just posterior to
the rim of the segment, due to the lateral margins bulging slightly
at this point; the apical margin of the flap deeply, roundingly
notched. Eleventh segment with an elongate fingerlike stylus as
in drawing.

The harpago, as viewed from a flattened position on a microscope
slide, is peculiar in shape in that its length is not much greater than
its width across its apex where its posterior dorsal corner is broadly
extended dorsad. At this dorsal posterior corner is located a small
external, sharply pointed hook.

The aedeagal structure has a more complex theca than in most
species. The aedeagus itself is extremely slender, showing only at
apical end beyond the theca, where it appears as a slender, goose-
necked structure which bends out from the theca and then back in
almost a complete circle, thus bringing the extreme apex under the

512 The University Science Bulletin

theca again. On the left side two sclerotized processes show, one a
long, flat, bladelike structure attached approximately at middle and
ending just cephalad of apex of thecal flap, the other a short,
sharply pointed hook which is only barely visible beyond posterior
margin of theca. On the right side only the tip end of an aedeagal
hook shows beyond the theca. The theca is a long semisclerotized
tube extending half way over the aedeagus. On the left side it ends
truncately at the middle, except at the dorsal corner, where it is ex-
tended caudad for some distance as a flap whose edges partially fold
together. On the right side the theca continues caudad as a broad
lobe which ends truncately at base of apical fourth of aedeagus.

Comparative notes. N. macidatus is readily distinguished from
all other species by the color of the tegmen, w'hich is translucent,
with one anterior oblique dark band and a partial posterior one, and
the color of the frons, which has two lateral dark longitudinal
streaks on each side of the median carina.

It more closely resembles N. perlucidus n. sp. and .V. vitripennis
Stal. From the former it differs by the presence of rudimentary
hind wings instead of long ones, by having a deep notch in pronotum
just behind eye, by a more wedge-shaped tegmen than a hemispheri-
cal one as in perlucidus and by having the combined distal part of
Cuia and CUj,, long, following the apical margin and thus setting
off a narrow phlange.

For comparison with A^. vitripennis Stal, see notes under this
heading in the description of that species.

Notes on distribution. Type locality is given as St. Cruz and
Sonoma, California. A few specimens have been collected by R. H.
Beamer in July, from the following places in California: Maria
county, Leona Heights, Niles, Upper Lake, Mt. Diablo, and Boulder
Creek.

Location of types. In the United States National Museum at
Washington, D. C.

Neaethus vitripennis Stal, 1854

(Plates LX, LXI, LXII, LXIII)
Stal, Carolus. Of. Vet. Aked. Forh., XI, p. 247. (Hysteropterum.)

Size. Length from apex of head to tip of tegmen is 4.4 mm. to
5.2 mm. Length of tegmen, 4 mm. to 4.4 mm. ; width of tegmen, 2.08
mm. to 4 mm.

Color. Entirely golden yellow. Tegmina transparent and all yel-
low. Eyes brown. Claws on tarsus black-tipped.

Structural details. Vertex broad and extremely short through

Doering: The Subfamily Issinae 513

middle where it is slightly more than one fifth of the anterior margin,
moderately produced beyond eyes, anterior margin slightly rounded.
Frons with lateral margins subparallel, outwardly rounding some-
what so that greatest width is across apical fifth, lateral margins
greatly elevated, median carina present down entire length. Post-
clypeus comparatively much longer than in N. maculatus or A'.
perlucidus. In this species its greatest length is equal to one lateral
margin of the frons. Pronotum much narrower than the head,
deeply notched behind eyes as in A^. maculatus, but with a thicker
lateral arm than in perlucidus, its anterior margin roundingly pro-
duced as an even hemisphere, its posterior margin shallowly convex.
Mesonotum through middle twice longer than pronotum, a median
longitudinal carina indicated and a short, lateral one, on each side
just mesad of each lateral bulla; a deep groove indicated just
posterior to the anterior margin. Tegmina less semicircular in out-
line with the costal margin more parallel to the claval margin than
in perlucidus; cells of wing large, angular and a good many less
cross veins than in other species; vein Sc^ straight and widely sep-
arated from the costal margin making a widely expanded costal
area, traversed by 8 to 9 veinlets and with the cells thus set off
wider through middle than at either end; vein M four-branched;
vein Cu branching considerably before middle and veins Cu^a and
Cujb thus separated by a longer cell than in maculatxis and others,
but united at apex and running as a combined vein just inside of
apical margin for a considerable distance before joining vein M^ and
with the latter vein setting off a narrow phlange for half the width
of the apical margin. Hind wings rudimentary, folded once longi-
tudinally.

Male genitalia. Anal flap (10th abdominal segment) a large,,
broad, flaplike structure, tubular at base and with its ventral, pos-
terior margin extended posteriorly, but to much less degree than in
maculatus, as a flaplike extension which is deeply and broadly
emarginate, so that it gives the structure a two-pronged effect.
Eleventh segment a small but distinct tube with a conspicuous
fingerlike stylus.

Each harpago, as viewed from a flattened view, is a subquad-
rangular plate with its posterior dorsal corner extended dorsad into
a triangular, sharply poiiited projection, and bears a small, ventrad-
pointing external hook located half way on the dorsal extension.

The aedeagal structure is complex. The aedeagus proper is an
elongate, tubular, well-sclerotized structure, which extends beyond

33_^141

514 The University Science Bulletin

the theca in goose-necked fashion and finally tapers to a sharply-
pointed apex which is bent back toward itself. From a left view it
shows four well-sclerotized hooks located as follows: one short,
sharply pointed one, located approximately at base of apical fourth;
another located approximately at middle of aedeagus and which
makes a semicircular curve ventrad after emerging from the thecal
rim ; a third and very small, sharply pointed lateral hook, located
just posterior to middle; and a fourth long, stiletto-type flap located
just above the lateral posterior hook. On the right side no new hooks
show, only the apices of the dorsal and ventral left hooks showing
beyond the thecal margin. The theca on the left side completely
covers the aedeagus only at base, then along dorsal side only ex-
tends caudad as a broad lobe to base of apical third of aedeagus,
where it is notched and forms two pointed prongs. On the right side
the theca covers the aedeagus all but its apical third or fourth.

Comparative notes. This species closely resembles the following
species: A^. maculatus Melichar and A^. perlucidus n. sp., in size,
translucency, in texture, the large angulate cells and the widely ex-
panded costal area. These three species are separated from each
other in the following ways: pronotum deeply notched behind eyes
in vitri'pennis and maculatus, but not so deeply in perlucidus ; teg-
mina held vertically to body in vitripennis and perlucidus, but much
more porrect in maculatus ; hind wings long in perlucidus, rudimen-
tary in other two; clypeus much longer proportionally to frons in
vitripennis than in perlucidus or maculatus; costal margin and claval
margin subparallel in vitripennis and maculatus, but distinctly out-
wardly rounding in perlucidus. V'eins Cuja and Cuii, united for only
a short distance at apex in perlucidus, with no narrowed phlange
set off along posterior border, while in vitripennis they are combined
for a longer distance and together with vein M^ sets off a phlange for
about one half the width of the apical margin and in maculatus for
about one third the width of the tegmcn.

Vitripennis Stal may very easily be confused with sinehamatus,
which in many respects it more closely resembles than any of the
above. For comparison, see the notes under this heading in the de-
scription of that species.

It also closely resembles superficially A^. siynilis n. sp. For com-
parison of this species see notes under this heading in the description
of that species.

Notes on distribution. In the National Museum at Washington,
D. C, are examples from Tamalpais and San Francisco, Cal. The
type locality is San Francisco.

Doering: The Subfamily Issinae 515

Neaethus sinehamatm n. sp.

(Plates LX, LXI, LXII, LXIII)
ORIGINAL DESCRIPTION

Size. Length from tip of head to apex of tegmen, 4.6 mm. to 4.8
mm. Length of each tegmen, 3.9 mm. to 4 mm.; width of each
tegmen, 2.4 mm.

Color. Like vitripennis and perluddus in being a uniform pale
golden yellow. Eyes grayish-brown. Veins at apex of clavus tinted
dark brown. Clavola of antenna blackish-brown. Tarsal segments
reddish-brown. Tip of claws on legs black.

Structural details. Shape of head and pronotum similar to per-
lucidus n. sp. Vertex twice wider than length at sides, its anterior
margin slightly angulate at middle, not produced beyond anterior
margin of eye, lateral margins parallel. Greatest width of eyes half
width of vertex. Frons and clypeus equal in length. Lateral margins
of frons elevated and diverging from posterior margin so that great-
est width is at base of apical fifth, a distinct median carina present
and disk concave on either side. Pronotum much narrower than
head, its anterior margin roundingly produced into vertex, but not
notched behind eyes, the lateral arm set off at this point much
shorter in length than in some species. Mesonotum one third wider
than length at middle, only a faint median carina indicated, each
lateral third buUate, a deep crescent-shaped groove following and
very close to posterior line of pronotum. Tegmen not semihemi-
spherical or parallel margined, but with greatest width in line with
apex of clavus. Veins of tegmen heavy, cross veins few and forming
large angulate cells. Vein Scj much closer to margin than in related
species, thus setting off a much narrower expanded costal cell than
in other species which is not recurved and traversed by only six or
seven cross veins; vein M only four-branched; vein Cui branching
just before middle of tegmen, then veins Cuja and Cuj,, again uniting
at apex and extending as a short ventrad-curving vein to meet M^.
Hind wings rudimentary, folded once longitudinally.

Male genitalia. Anal flap (10th abdominal segment) much shorter
than in vitripennis Stal. Its expanded ventral apical margin not any
longer than the tubular basal part, the extreme apex very shallowly
emarginate. Eleventh segment only slightly visible and bearing a
short stylus.

Each harpago, as viewed from a flattened view, is a subquad-
rangular plate with its posterior or distal end extended dorsad as a
projection which is slightly bulbous at base where it bears an ex-

516 The University Science Bulletin

ternal, ventrad-directed pointed hook and then ends in a spatulate
lobe.

The aedeagus on the left side appears as a semimembranous
tubular organ bearing several well-sclerotized processes located as
follows: one attached at extreme base, gently curved in a semicircle,
and ending in a point just beyond middle; a second long, fiat process
attached at middle is parallel-margined for three fourths of its own
length and has its apical bent end ending in a fine point not far
distant from apex of aedeagus; a third very short spine attached just
anterior to middle. On the right side no additional hooks of the
aedeagus are visible. The theca is unusual in that it covers only
the extreme base of the aedeagus on the left side, but on the right
side is extended into a long spatulate lobe which covers all of the
aedeagus except the extreme apex. See drawings.

Comparative notes. This species more closely resembles vitri-
pennis Stal and perlucidus n. sp. than any others because of its same
coloring, size, etc. It is separated from perlucidus by having rudi-
mentary hind wings, while perlucidus has long ones; by having the
anterior margin of vertex not produced beyond eyes, by having more
wedge-shaped tegmina which are wider in line with apex of clavus,
while in the former they are hemispherical, and finally by having a
very narrow costal cell area which is not bent back as in perlucidus.

From vitripennis Stal it is distinguished by not having the vertex
so produced anteriorly or not nearly as wide, by not having the an-
terior margin of pronotum notched as in that species, and by having
the costal cell area much narrower, not bent back and with fewer
cross veins in it. In addition, male specimens can easily be sep-
arated by the external appearance of the anal flap, which is much
larger and two-pronged in intripennis, but is only shallowly emargin-
ate in sinehamatus.

Location of types and type locality. Described from fourteen
specimens. Holotype, male, and allotype female taken at Apline,
Cal., August 9, 1929, by Dr. R. H. Beamcr. Three paratype males
from same locality. One paratype female from Campo, Cal., August
10, 1935; one paratype male and seven paratype females from San
Diego county, California, August 7, 1929, collected by Dr. R. H.
Beamer. The types are in the Snow Entomological Collection, Uni-
versity of Kansas.

Doering: The Subfamily Issinae 517

Neaethus similis n. sp.

(Plates LX, LXI, LXII, LXV)
ORIGINAL DESCRIPTION

Size. Length of body from apex of head to tip of tegmen, 3.9 mm.
to 4.9 mm. Length of tegmen, 3.2 mm. to 4.6 mm.; greatest width
of tegmen, 3.2 mm. to 4.6 mm.

Color. General color similar to N. fenestratus. Body reddish-tan.
Vertex and pronotum margined in dark brown, a faint light streak
indicated on median line; two round, reddish-brown depressed spots
in middle of pronotmn. Mesonotmn yellowish-tan through middle,
darker at sides. Eyes yellowish-tan, banded with red bands. Frons
uniformly dark tan, disk on either side of median carina sometimes
darkened, margins dark brown. Body dark or golden-tan, the edges
of some of the segments dark brown, and in females the median
border of the ovipositor valves blackish-brown; males more uni-
formly colored. Legs reddish-tan, sometimes washed in darker brown
on carinae or the ends of segments; tips of the claws black. Tegmina
amber, translucent, sometimes somewhat milky, but not as much so
as in N. jacintiensus. Veins uniformly dark brown in females; in
male longitudinal ones sometimes lighter.

Structural details. Head not much wider than pronotum. Vertex
not as broad as in jacintiemus but narrower than N. vitripennis, be-
ing about four and one half times wider across anterior margin than
at median line, and being produced beyond eye for at least one
fourth length of lateral margin of vertex, the lateral margins of the
latter considerably diverging posteriorly. Frons long, narrow, par-
allel-margined with greatly elevated lateral edges. Postclypeus
comparatively short, about one fifth shorter than one lateral margin
of the frons. Pronotum not notched behind the eye, as in vitripennis,
its anterior margin considerably elevated and roundingly produced
as an even hemisphere, its posterior margin shallowly convex, the
lateral arms at side, behind eyes, about equal in length to one
fourth pronotum along median line; two round pits in center of the
disk. Mesonotum one third wider than length through middle, a
deep groove along anterior border forming an arcuated line just
posterior to margin of the pronotum ; somewhat shallowly depressed
through middle, leaving ,an elevated shoulder on each lateral fourth;
a median carina sometimes faintly indicated. Shape of tegmen not
as hemispherical as in many species, its greatest width in line with
apex of clavus. Veins not as heavy as in jacintiensus. Cross veins
fewer and parallel, thus making many parallel-sided cells, not so

518 The University Science Bulletin

much smaller at apex than elsewhere. Vein Sc^ straight or only
slightly curved outwardly, setting off a wide costal phlange, which is
distinctly inflexed and is traversed by seven to nine cross veins,
making practically all the cells at either end smaller than through
middle. Veins SCo and R usually w'ith two branches, sometimes
three; vein M pectinate, usually with six branches; vein Cu^
branched near base, as in vitripennis, and is united at apex, where it
bends ventrad to meet a branch of M, which results in a narrow
phlange being set off for about one third of the apical margin. Hind
wings rudimentary, folded once longitudinally.

Male genitalia. Anal flap (10th abdominal segment) of the male
makes the species easily recognized, since its posterior margin is
deeply emarginate, which gives it the appearance of being pronged.
The tubular part is slender and has its outer margins somewhat
bulging, so that its greatest distance is through middle. The eleventh
segment shows as a ringlike lobe beyond the dorsal posterior margin
of the tenth and bears a more triangular-shaped stylus than in many
species.

The harpago, as viewed from a flattened lateral view, is a boot-
shaped plate which is narrowed at the apex, then greatly bulged on
the ventral side through the middle, after which it narrows again to
a blunt, truncate apex, and at the base of this short projection bears
an external ventrad-curving, slender hook.

The aedeagal structure in this species is complex, especially as
regards the theca, which covers the aedeagus for most of its length.
The apical portion of the aedeagus, which shows beyond the theca.
is membranous and therefore wrinkled on its inner margin. On the
left side the theca appears as a membranous tube with its opening
about halfway from base of aedeagus. At this point the tips of two
well-sclerotized hooks of the aedeagus show beyond the emarginate
outer margin of the tube, the dorsal one a fiat, sharply pointed proc-
ess, attached somewhat near base, and the ventral one a shorter
bifurcate process. The inner margin of the thecal tube extends
caudad as a flat membranous lobe, whose posterior margin is
slightly triangularly produced and beyond which the apex of a third
fiat, sharply pointed aedeagal process shows, which is attached near
middle of aedeagus. On the right side the theca shows as a longer
membranous tube with an opening near the apex and beyond which
shows a sharply pointed, sclerotized hook of the aedeagus. From a
microscope slide it is difficult to understand the twisting of the
theca. Apparently the usual dorsal flap is split dorsad and the

Doering: The Subfamily Issinae 519

lateral lobe has twisted around the acdeagus from a point midway on
the aedeagus so that it makes it appear to have two openings.

Comparative notes. Neaethus similis n. sp. is very similar in
superficial appearance to vitripennis and jacintiensus n. sp. It has,
however, one of the most distinctive types of genitalia in the genus.
These three species can be separated externally by the following
characteristics: the anterior margin of the vertex is produced be-
yond eye for about one fourth the length of its lateral margin in
similis, wiiile in the other two is scarcely produced at all and the
width across this margin is less in sijyiilis, being only about four and
one half times the length on median line, while in vitripennis it is five
times, and in jacintiensus five and one half. Similis has a longer and
narrower frons and is about one fifth shorter at lateral margins than
greatest length of clypeus; jacintiensus has the frons about eciual
in length and width, while vitripennis has a longer clypeus propor-
tionally, its length being equal to one lateral margin of the frons.

The anterior margin of the pronotum is notched behind the eyes
in vitripennis, but not in the other two. The tegmina show a few
variations, the costal cell area is reflexed distinctly in sirnilis and
vitripennis but not in jacintiensus ; vein M tends to branch more in
jacintiensus, showing usually as many as seven branches, while
similis frequently has six, and vitripennis only four. The anal flap
of the male is pronged in similis and vitripennis and rounded in
jacintiensus.

Location of types. Described from holotype male collected at
Leona Heights, Cal., August 15, 1933; allotype female, Mt. Diablo,
Cal., August 21, 1935; one paratype female, Mt. Diablo, Cal., and
four paratype males from Mt. Diablo, Cal., collected by Dr. R. H.
Beamer; one paratype female and seven males from Mt. Diablo,
Cal., collected by P. Oman. These types are in the Snow Entomo-
logical Museum, University of Kansas, and the U. S. National Mu-
seum at Washington, D. C.

Neaethus jacintiensus n. sp.

(Plates LX, LXI, LXII, LXV)
ORIGINAL DESCRIPTION

Size. Length from tip of head to apex of tegmen is 4.3 mm. to 5.6
ram. Length of tegmen is 3.8 mm. to 4.8 mm.; width of tegmen is
2.4 mm. to 3.1 mm.

This is one of the largest species in the genus, approximating
N. jragosus in size.

Color. General color light yellow washed in varying shades of

520 The University Science Bulletin

fuscous so that some specimens appear much darker than others.
Vertex light brown to fuscous with broad, light-yellow margin and
a median longitudinal band light. Eyes mottled brown. Prothorax
light brown to dark except at margins and a median longitudinal
line. Mesonotum light brown to reddish across the depressed middle
portion, lateral bullae dark brown, the anterior margin and apex
light yellow. Frons and clypeus uniform, in the lighter specimens
being light-tan or yellow, with the clypeus slightly lighter; in the
darker specim?ns being reddish-brown with the margins light yellow.
The underside of body light yellow with touches of light green,
especially on the third and fourth segments, or most of segments
having the central parts of sclerites reddish-brown with margins
light yellow and the abdomen dark brown on the third to sixth seg-
ments. Legs light yellow or reddish-brown, tips of claws black.
Tegmina milky translucent with either dark-brown veins entirely
or some of the longitudinal veins lighter.

Structural characteristics. Head not much wider than pronotum.
Vertex broad, its width across anterior margin about five and one
half times its length at middle, the anterolateral corners not any
produced beyond the eyes, but the anterior margin slightly angu-
lately produced at middle, the lateral margins distinctly diverging
posteriorly. Pronotum four times wider than length at middle,
gradually narrowed behind eye to a slender arm, but not at all
notched. Mesonotum broadly triangular, much more than one
third broader than long, depressed through middle with a median
carina lacking or only slightly indicated. Length and width of
frons equal, its lateral margins outwardly rounding and slightly
elevated, a median carina present for three fourths its length, on
each side of which the disk is depressed as far as the elevated lateral
margins. Clypeus not as inflated as in some species, about one fifth
shorter than greatest lateral margin of frons. Tegmen with greatest
width about two thirds its length and at a point in line with apex
of clavus and not in middle of wing as in nigronervosus or fenes-
trates; veins thick, many cross veins present, thus making many
angulate cells, the cells at the apex short. Vein Scj straight and
long, extending to a point which is at least one third length of
tegmen, the cells in the costal area uniform, rectangular cells. Veins
Scg and R usually each only two-branched; vein M pectinate with as
many as seven branches as in nigronervosus; vein Cu branched
nearer base than in other similar species, united again close to apex.
Hind wings rudimentary, folded once longitudinally.

Doering: The Subfamily Issinae 521

]\[ale genitalia. Anal flap (10th abdominal segment) a broad, flat
tube at base with its lateral basal margins parallel-sided, its ventral
posterior margin extended as a short, broad flap with rounded apex.
From a mounted specimen the eleventh segment is exposed as a ring-
like segment, showing beyond the dorsal rim of the tube, and bears
a fingerlike stylus which is as long as the apex of the flap.

From a flattened view each harpago appears as a flat, somewhat
inflated lobe, which is broadest at about middle, where its ventral
margin is outwardly rounded. At the posterior dorsal angle the
harpago is expanded into a truncate flattened process which is
abruptly bent ventrad and posteriorly on the inside, and on the out-
side bears a sharply pointed but short hook whose apex is directed
directly ventrad.

The aedeagus, as viewed from the left side, appears as a slender,
semicircularly curved tube, bearing two well-sclerotized processes,
the basal one being much shorter, attached at base under the theca
and ending about midway of aedeagus, only its apical part showing
beyond theca as a curved hook, which is broader at base, then tapers
to a sharp apex which gently curves ventrad. The other aedeagal
process arises midway on aedeagus as a broad, knifelike structure,
and gradually tapers to a fine point, which stops approximately at
base of apical ninth of the aedeagus. On the right side the aedeagus
is mostly covered by the theca. It shows a flat, bladelike process at-
tached near base and ending at about the base of the apical third.
The theca on the left side covers the aedeagus for about one fourth
its length, and its posterior margin on this side is deeply emarginate.
On the right side the theca extends tubelike to not quite the middle
of the aedeagus, then has a dorsal flaplike extension which reaches
almost to the tip of the aedeagus and is sharply pointed.

Comparative notes. N. jacintiensus is closely allied to A^. ni-
gronervosus Mel. and A', fenestratus Mel. It is separated from
these species by several characteristics. It is much larger than either
of the other two. The vertex is proportionally much wider and the
length at middle much shorter; the posterior lateral margins dis-
tinctly diverge. The pronotum also is proportionally wider for its
length. The mesonotum lacks a median carina which the other two
usually have indicated. The tegmen in this species is not hemi-
spherical and has its greatest width in line with apex of clavus in-
stead of at middle. The cells of the tegmen near apex are short,
not long and parallel-sided, as in nigronervosus, nor clustered at apex,
as in fenestratus. The venation resembles that of nigronervosus

522 The University Science Bulletin

more than it does fenestratus, as M tends to be pectinate with many-
distinct branches.

N. jacintiensus n. sp. also resembles A'', similis n. sp. For compari-
son of these two species see notes under this heading in the descrip-
tion of that species.

Notes on distribution. A large series of this species was taken in
San Jacinto mountains, California, by Dr. R. H. Beamer. Other
specimens have been taken at the following places in California:
Campo, Big Bear Lake, Idyllwild, San Diego county, Giant Forest,
and Beaumont.

Location of types. Holotype male, and allotype female collected
in the San Jacinto mountains, California, by R. H. Beamer, August
21, 1929. Many female and male paratypes taken at the same time
and the same places as listed above. The types are in the Snow
Entomological Collection, University of Kansas.

Neaethus nigronervosus Melichar, 1906

(Plates LX, LXI, LXII, LXV)

Melichar, Leopold. Monogiaphie der Issiden (Homoptera). Alili. k. k. Zool.-Bot. Ges.
Wit-n, III, pt. 4, 1906.

Size. Length of body from apex of head to tip of tegmen, 3.2 mm.
to 4.5 mm. Length of tegmen, 2.6 mm. to 3.7 mm.; width of tegmen.
1.5 mm. to 2.4 mm.

This species is one of the smaller species in the genus.

Color. This species generally darker than many in the genus.
Vertex fuscous or somewhat dark tan through middle, margins and
a median longitudinal band light yellow. Pronotum also fuscous or
darkish all except margins and median light line. Mesonotum gen-
erally brown except light yellow on anterior border at apex, median
carina and on lateral fourth except for two brown spots, one at ex-
treme anterior, lateral corner and another in center of the bulla.
Eyes variegated with brown. Frons all dark brown except light in
the following places; all the outside margins, the median longitudinal
carina, which forks slightly at apex so that the yellow spreads out
into a spot on each side, and on each side a narrow, irregular longi-
tudinal streak just mesad of each lateral margin. Genae with a
brown streak anterior to eye, rest light yellow. Antenna yellow,
washed in brown, with a dark brown clavola. Clypeus dark brown
except for a median band down center and at each lateral carina.
Prothorax and mesothorax on underside brown, margined in whitish
yellow. Metathorax light. Abdomen of female blackish-brown ex-
cept for a light spot in center of first few abdominal segments, an-

Doering: The Subfamily Issinae 523

other at lateral corners of or entirely across the sixth and seventh
abdominal sterna, all of the abdominal plem-a, and the central por-
tions of the ovipositor valves. Males with venter of thorax light
yellow spotted with dark; abdomen yellow except the fifth, sixth,
and seventh abdominal segments blackish brown. Legs brown except
for yellow along the carinae and tip of claws, which are black.
Tegraina milky translucent with dark brown to black veins.

Color variations. Melichar distinguished this species from N.
fenestratus by stating that all the veins are black in nigronervosiis,
while in fenestratus the longitudinal veins are greenish and only the
cross veins are blackish. In all the males of nigronervosus studied
by the writer the longitudinal veins are always lighter than the
transverse veins.

Furthermore, some males have the entire body light yellow washed
in lighter reddish-brown.

Structural details. Vertex not deeply concave through middle nor
produced anteriorly beyond eyes, the anterior margin straight or
only gently rounding, its width across anterior margin four times its
length at middle. Frons rectangular, the lateral margins only
slightly converging at base, somewhat elevated and a median carina
present, which lessens in height and finally disappears as it ap-
proaches the apex. Postclypeus as long as or longer than the lateral
margin of frons. Pronotum broad, about four and one-half times its
length at middle, its anterior margin rounding, but not much pro-
duced into vertex, narrowed behind eye to a thin arm, but not deeply
notched in this region as in other species, two depressed pits in center
of disk. Mesonotum concave through middle, raised on either side
to a distinct bulla; a median carina present, and an arcuated groove
at extreme anterior margin following the posterior margin of the
pronotum but stopping at either bulla. Tegmina hemispherical in
outline, its greatest width just anterior to apex of vertex; its length
approximately one half greater than its width; veins thick but ap-
pearing somewhat thinner than fenestratus, many cross veins pres-
ent, forming numerous, irregularly shaped cells, with those at apex
long, narrow, and parallel-sided, vein Sc^ widely separated from
costal margin, arcuated inwardly rather than straight or the reverse
and only extending posteriorly for a third or less of the wing; the
other longitudinal veins not uniform as to their branches, frequently
pectinate to the extent that Sc^ may have three to four branches and
M frecjuently five to seven ; Cu usually only two branched and these
branches frequently making a curve ventrad but not uniting. Hind
wings reduced to a mere scale, without longitudinal fold.

524 The University Science Bulletin

Male genitalia. Anal flap (10th abdominal segment) short, not
much more than a third longer than wide, the ventral posterior
margin extending flaplike for less than half the total length, evenly
rounding at apex. The eleventh segment more conspicuous than in
some species, with a long, fingerlike stylus extending beyond the
apex of flap.

Each harpago, as viewed from a flattened view, is a broad, inflated
lobe whose ventral margin is strongly outwardly convex and whose
dorsal margin is deeply concave at middle. The posterior dorsal
corner is projected dorsad into a bifurcate process, the more dorsal
hook slenderly pointed and abruptly recurved, the ventral hook di-
rected caudad and bluntly pointed.

The aedeagus is a semisclerotized tube, the apical fourth of which
is strongly recurved and somewhat narrowed. From a left view it
shows two strongly sclerotized, flat, sharply pointed processes which
are attached some place near base and extend caudad, one having a
more dorsal position and following the curve in the aedeagus until it
reaches the apical bend in the latter and the other laterad in posi-
tion and extending for about the same distance caudad, but with its
tip abruptly recurved. On the right side the aedeagus appears to
show a similar lateral process of about equal size and shape as the
one on the left side and the dorsal process shows from this view, too.
The theca in this species is a simple semimembranous tube with its
posterior margin on either side obliquely truncate.

Comparative notes. This species is one of the most variable
species in the genus in size, coloring and certain structural details.
Melichar apparently had only females on which to base his de-
scription. No males in any collection seem to fit his description of
the veins which he stated were all black. The males seem to have
lighter longitudinal veins, as in fenestratus.

N. nigr-onervosus resembles N. fenestratus and jacintiensus more
than any other species. Nigronervosus and fenestratus are separated
other than the male genitalia, which are vastly different, by only the
following characteristics: the vertex in nigronervosus has a straight,
or only slightly rounding anterior margin, while in fenestratus this
margin is slightly angulate at middle; the costal cell area is ex-
tremely broad in nigronervosus due to the fact that vein Sc^ bends
inwardly instead of toward the costal margin as in the other species,
where the area is quite narrow ; the cells along the apex of the tegmen
in nigronervosus are long, narrow and parallel-margined, while in
fenestratus there is always a cluster of small cells at apex. This
latter is the most distinctive thing between the two.

Doering: The Sibfamily Issinae 525

In addition to the above there are trends of differences in the wing
venation which can be noted. These differences are not always ap-
parent because there seems to be some variation in the species
themselves due to the longitudinal veins breaking up into various
patterns of reticulation. However, there is a marked tendency for
vein M in nigronervosus to be pectinate, with from six to seven
branches, and for these veins to bend down and encroach on the terri-
tory of the R vein, while in fenestrahis M is usually only four-
branched, its branches are usually in the dorsal half of the corium
and the veins Scj and R are more apt to be pectinate.

For comparison with jacintiensus see the description of the latter.

Notes on distrihutioyi. The type locality is given as Prescott,
Ariz. Many specimens were studied from the following places in
Arizona: Yarnell, Chiricahua mountains, Granite Dell, Kirkland
Junction and Maricopa county. A few specimens from Cedar City,
Utah, were available.

Location of types. Female holotypes in the National Museum,
Washington, D. C.

Neaethus jenestratus Melichar, 1906

(Plates LX, LXI, LXII. LXIII)

Melichar, Leopold. Monographie der Issiden (Homopteia). Abli. k. k. Zool.-Bot. Ges.
Wien, III, pt. 4, 1906.

Size. Length of body from tip of head to apex of tegmen, 3.4 mm.
to 4.2 mm. Length of tegmen, 2.9 mm. to 3.5 mm. ; width of tegmen,
1.8 mm. to 2.6 mm.

This species approximates nigronervosus in size and is one of the
smaller species in the genus.

Color. Melichar describes this species as having a pale green
body and with wing covers transparent and longitudinal veins pale
greenish, while the cross veins are entirely black. In a series of speci-
mens from several localities these color characteristics seem variable.
A female specimen from the National Museum Collection, bearing a
Melichar determination label, is as follows: vertex yellowish-tan
with the thin, sharp margins dark brown, pronotum yellowish-tan
with two round reddish pits midway of its length, extreme thin
margins dark brown; mesonotum yellowish-tan except for a light
reddish-brown spot at lateroanterior corner of the bulla and another
on the posterior half of each bulla. Frons uniform yellow with
dark-brown margins, postclypeus yellow except for six or seven
pairs of faintly indicated oblique stripes; rest of head uniform yel-
lowish-tan. Thorax and abdomen all yellowish-tan, except the

526 The University Science Bulletin

inner margins of each ventral valve of ovipositor. Legs yellow ex-
cept for tips of spines and claws. Tegmina milky translucent; longi-
tudinal veins light yellow or tan, cross veins dark brown.

Color variations. Specimens which the writer has placed under
this species show many color variations. Some specimens show all
the veins fairly light ; others show them all to be dark brown. When
the veins are all dark much fuscous is found on the rest of the body;
vertex, pronotum, and mesonotum are washed in it or show it at the
sides; on underside all the sclerites are mottled with it and the
fifth, sixth and part of the seventh segments are entirely fuscous in
the female.

Structural details. Vertex not produced beyond eyes, its anterior
margin slightly angulate at middle, its width at this point four times
the length of one lateral margin. Frons broad, approximately one
fourth longer than width across basal line, its lateral margins ele-
vated, several depressions or dimples on each lateral half, a median
carina present for three fourths of its length. Postclypeus longer
through middle than the lateral margin of the frons, a deep groove
present at point of union of the two sclerites. Pronotum short and
broad, not much narrower than head, its length at middle twice the
length of the vertex at middle, its length behind eye abbreviated to
a mere slender arm, the anterior margin not greatly notched out as in
some species, two round depressions present in the middle of the disk.
Tegmina hemispherical in outline, its length not quite one third
greater than its greatest width, which is midway of the wing or an-
terior to apex of vertex. Vein Scj short, not extending even as far
as one third the entire length, much nearer the costal margin and
not curved inwardly as in nigronervosus, the other longitudinal
veins frecjuently forked just inside apical margin and with many
cross veins forming a cluster of irregular small cells at apex of
corium. Vein Sc, frequently pectinate with four branches present,
vein R with two or three main branches, vein M is usually four-
branched, but its branches are all in dorsal half of corium and vein
Cuj has the usual two branches which approach each other just be-
fore apex, but do not unite, and due to the high position of the medial
veins curve for only a short distance at apex. Hind wings reduced
to a mere scale, without a longitudinal fold.

Male genitalia. Anal flap (10th abdominal segment) short, not
much over one fourth longer than wide, its lateral margins curving
outwardly, its ventral margin extended only a brief distance to form
the flap proper, the extreme posterior margin of which is truncate.

Doering: The Subfamily Issinae 527

The eleventh segment only sliglitly visible, its stylus moderately
long, extending not quite to apical margin of the stylus.

Each harpago, as viewed from a flattened lateral view, is a broad,
somewhat inflated lobe, whose ventral margin is strongly outwardly
convex and whose dorsal margin is deeply and angulately concave.
The posterior dorsal corner is projected dorsad into a triangular
process, which is broad at base and then tapers to a slender, recurved
apex, just ventrad of which is a small, recurved external hook.

The aedeagus is a semisclerotized tube, the apical fourth of which
is strongly recurved and bluntly rounded at apex. From a left
view it shows two distinct sclerotized processes, the most conspicuous
one located more ventrad, attached somewhere near the base under
the theca, then after emerging from the theca immediately dividing
into two hooks, a short dorsal one which is sharply pointed, half the
length of the other and points ventrad, the other one long, slender,
not quite so sharply pointed and curved dorsad, ending at about
the base of the apical fourth of the aedeagus; the second aedeagal
hook on this side is partially concealed by the bifurcate process and
shows only as a sharply pointed, flat blade between the dorsal short
hook of the latter and the right thecal flap. On the right side the
aedeagus has a flat, bladelike, sharply pointed process, lying closely
adpressed to it, attached midway the length of the aedeagus and
ending approximately at base of apical fourth. The theca is a semi-
sclerotized tube, covering the aedeagus for about one fourth to one.
third its length and on the dorsal side is extended caudad as a
broad, spatulate lobe which becomes slightly wider at its apex.

Comparative notes. This species resembles A^. nigronervosus and
A^. jadntiensus n. sp. more closely than any others. For comparison
with these species see notes under this heading in the descriptions of
the other species.

Notes on distribution. The type localities are given as Los
Angeles, St. Cruz, and Claremont, Cal.

A specimen in the National Museum collection, bearing a Meli-
cliar determination label, is from Arizona. Specimens were on hand
for study from the Chiricahua mountains. Congress Junction, Oak
Creek Canyon, Santa Rita mountains and Gila, Ariz. A few speci-
mens were studied from Silver City, N. Mex., and Cedar City, Utah.

Location of types. The types from Los Angeles and Santa Cruz,
Cal.. arc in the National Museum collection at Washington, D. C.
The Claremont, Cal., specimens, collected by Baker are in the
Melichar collection.

528 The University Science Bulletin

Neaethus curvaminis n. sp.

(Plate LX, LXI, LXII, LXIV)
ORIGINAL DESCRIPTION

Size. Length of body from apex of head to tip of tegmen, 3.4 mm.
to 4.1 mm. Length of tegmen, 2.8 mm. to 3.4 mm.; width of tegmen,
1.8 mm. to 2.3 mm.

This is one of the small species in the genus, approaching the size
of uniformus and diversus.

Color. Typical general color stramineous as in uniformus, some-
times veins darker as in the nigronervosus group. Vertex and prono-
tum stramineous, with thin outer edges dark brown and a cream-
colored median line. Eyes reddish-brown. Mesonotum dark tan
with an arcuated light border around anterior margin, which bends
posteriorly at either side and crosses each lateral bulla; the median
carina and extreme apex also cream-colored. Frons uniformly
stramineous with thin outer margins dark brown and median carina
lighter. Clypeus yellow with a series of oblique brown bars on each
side. Under and lateral sides of thorax and external genitalia golden
yellow, abdominal segments cream or light green. Inner margins of
ovipositor valve dark brown. Legs washed in reddish-brown; tips
of claws black. Tegmina stramineous throughout, usually with the
veins the same color.

Color variations. Some specimens have the frons, vertex, prono-
■tum, and mesonotum infuscated to varying degrees and the veins
of tegmina dark brown.

Structural details. Similar in appearance and structure to uni-
formus and diversus. Vertex narrow, being only twice as wide across
its anterior border as a lateral margin or approximately twice as
wide as eye, this anterior border rounding, and posterior margin
abruptly elevated. Frons elongate, parallel-sided, not much nar-
rowed across posterior or basal margin, lateral margins somewhat
elevated, median carina not greatly pronounced, extending only three
fourths of the total length. Postclypeus long, being as long as or a
trifle longer than the lateral margin of the frons. Pronotum longer
than in related species, being three times wider than greatest length,
narrowed in the characteristic fashion behind the eyes to a slender,
somewhat tapering lateral arm, but not notched at this point.
Mesonotum broad, about twice as wide as long, an arcuated groove
following the margin of the pronotum and a faint median carina
indicated. Tegmina hemispherical in shape with costal and apical
borders strongly rounding, their greatest width at middle of the

Doering: The Subfamily Issinae 529

wing, clavu:^ moderate in length, being only twice longer than dis-
tance beyond its apex. Vein Sc^ moderately short, stopping ap-
proximately at apex of basal fourth of tegmen, inwardly arching so
that the costal cell area is broad and being crossed by only four to
six cross veins, making large, distinct cells which are always broader
than long. In this species the other longitudinal veins usually have
one extra branch, the characteristic pattern being SCo eitiier two- or
three-branched, R three-branched, M five-branched with all its
branches distinctly in the upper half of the wing, Cuja and Cu^b
combined before apex, but the combined vein not uniting with an M
vein. Hind wings reduced to mere scales.

Male genitalia. Anal flap (10th abdominal segment) short, nar-
rowed sliglitly at base, bulging through middle and ending on the
ventral side in a truncate flap. The ringlike eleventh segment shows
beyond the dorsal posterior margin and bears an elongate stylus
whose apex extends beyond the tip of the flap.

Each harpago, as viewed from a flattened lateral view, appears as
a pear-shaped plate, somewhat inflated through middle. At the
posterior corner it is reflexed so that an internal phlange is formed
for about one fourth its length. The outer posterior dorsal corner
of the harpago is extended dorsad into a bluntly pointed hook and
the extreme posterior inward corner of the phlange extends ventrad
as a sharply pointed hook, the two liooks together making a
bifurcate corner to this region of the harpago.

The aedeagus is a short semimembranous tube, which is somewhat
broadened and bent forward at the extreme tip. On the left side it
shows tln-ee processes, the longest being ventral in position, more
heavily sclerotized and attached somewhat near the base of the
aedeagus. It is strongly curved, tapers at the apex and ends at the
point where the aedeagus curves cephalad. Another well-sclerotized
but shorter hook is seen just above the former and extends only to
about the middle of the aedeagus. Between these well-sclerotized
hooks is a third, elongate semimembranous lobe which is only
slightly longer than the dorsal hook. On the right side the aedeagus
bears a sharply pointed, curved hook attached at apex of basal third
and extending to base of apical third.

The theca in this species, as in vniformus, is conspicuous. It
covers the aedeagus for 'a little over one fourth of its length as a
tight-fitting tubular sheath. On the left side it ends truncately. On
the right side its dorsal corner is extended caudad into a flat, tri-
angularly-shaped flap, which extends slightly beyond the apex of the
lateral aed'^agal hook.
34—4141

530 The University Science Bulletin

Comparative notes. This species is closely allied witii A^. uni-
formus and A^. diversus. For comparison of the three species see
notes under this heading in the description of A^. diversus.

Notes on distribution and location of types. The species was de-
scribed from numerous specimens taken at various places in Arizona.

Holotype male, collected July 10, 1933. and allotype female,
August 14, 1935, at Granite Dell, Ariz., by R. H. Beamer. Five
paratype males and five paratype females, also from Granite Dell,
one paratype male and two paratype females from Santa Rita moun-
tains, four paratype males and six paratype females from Oak Creek
Canyon, one paratype male and two paratype females from Yarnell,
two paratype males from Prescott, one paratype male, Gila, and
one paratype each sex, Congress Junction, all collected by R. H.
Beamer in July and August.

These types are in the Snow Entomological Collection at the Uni-
versity of Kansas.

Neaethus uniformus n. sp.

(Plate LX, LXI, LXII, LXIV)

original description

Size. Length of body from tip of head to apex of tegmen, 3.6 mm.
to 4.2 mm. Length of tegmen, 3.1 mm. to 3.6 mm.; width of tegmen,
1.8 mm. to 2 mm.

Color. Similar in color to A^ diversus. General color, amber yel-
low. Vertex lighter cream yellow at posterior lateral corners, its
anterior margin dark brown. Pronotum slightly darker than vertex,
its margins also brown. Mesonotum like pronotum except for a light
cream anterior border anterior to the arcuated groove, which con-
tinues as a light streak across the middle of each lateral bulla, ending
at the sides as a spot ; the median carina and median posterior mar-
gins also cream. Eyes spotted and ringed with reddish-brown. Most
of frons amber, light cream along lateral borders and the thin outer
margins dark brown. Clypeus amber with a broad median cream
longitudinal band. Legs light yellow to amber, with carinae of
femora cream and tips of tarsal claws black. Thorax and abdomen
in both sexes cream. Ventral valves of ovipositor along the median
line dark brown. Tegmina semiopaque light amber; all of veins
cream except cross veins in cell Cu^b and apex of clavus, which are
sometimes dark brown.

Structural details. Vertex not produced much beyond eye, the
anterior margin slightly rounding; four times wider at its anterior
margin than its median length and twice the width of one eye; hind

Doering: The Subfamily Issinae 531

margin abruptly elevated. Frons short, clypeus long, the greatest
length of latter equalling the length of one lateral margin. Pronotum
three and one half times the greatest length, narrowed behind eye
but not notched, to form a slender, somewhat tapered lateral arm;
disk depressed through middle with two round, faintly depressed
spots in exact center. Mesonotum one third wider than long, a
median carina present and an arcuated groove close to anterior
margin, which ends at either side in a bulla. Tegmina more hemi-
spherical in outline than in diversus, their greatest width approxi-
matel}' at middle; vein Sc^ quite straight, and costal cell area only
moderately wide, with six, seven or eight cross veins, so that the
small cells in this area are usually square or approximately as wide
as long. The longitudinal veins coarse in texture with characteristic
branching for the genus ; veins Cu^a and Cu^b either not united at all
before apex or lost in the apical reticulation. Hind wings reduced
to mere scales.

Male genitalia. Anal flap (10th abdominal segment) short, nar-
rowed slightly at base, then broadening posteriorly. The ventral
posterior edge of the tube extended into a rounded lobelike flap which
is shorter than in most species but is a trifle longer than in diversus.
The stylus of the eleventh abdominal segment is blunt and short.

Each harpago, as viewed from a flattened lateral view, appears as
a pear-shaped plate, somewhat inflated or swollen through middle.
At the posterior corner it is reflexed for about one third of its dorsal
length. The extreme posterior edge of this refiexed portion is ex-
tended into two short, sharply pointed processes, one of which ex-
tends inwardly and dorsad and one of which extends caudad and ex-
ternally.

The aedeagus is a short, semimembranous tube, the tip of which
curves dorsad in a semicircular position. On the left side it shows
three sclerotized hooks or lobes. The most heavily sclerotized of
these is attached under the theca, somewhat near the base. Beyond
the rim of the theca it shows as a bifurcate process bearing a pointed
blade which extends to a point at base of apical third of aedeagus
and a small, sharply pointed hook which projects inwardly and is
usually partly concealed by the thecal hood. Another heavily
sclerotized hook is attached near base of apical third and extends
to about base of apical fourth. Just dorsad of this hook is a less-
sclerotized lobate-process, which ends at about the same point as the
latter. The theca in this species is conspicuous. It covers the
aedeagus for about one third its length as a tight-fitting tubular
sheath. Beyond this point the dorsal portion becomes expanded

532 The University Science Bulletin

into a large hood which from either side view is rather triangular in
outline. In reality it is a plate which has been elevated along its
middorsal line into a thickened ridge and has each lateral margin
extended downward as a lobate flap which partially encloses the
aedeagal hooks.

Comparative notes. This species is closely allied to the following
species: A^^. diversus and A^. xinijormus. For comparison of these
three species see notes under this heading in the description of .V.
diversus.

Notes on distribution and location of types. This species was de-
scribed from a large series of specimens taken in the Santa Rita
mountains, Arizona. Holotype, male, Santa Rita mountains, Ari-
zona, August 17, 1932; allotype female same place, August 18, 1935,
by Dr. R. H. Beamer. Eight paratype males and five paratype
females were collected same place in July and August, by Dr. R. H.
Beamer. Forty male paratypes and thirty-four female paratypes
were collected same place by F. H. Snow, in June. All types in the
Snow Entomological Collection, University of Kansas.

Neaethus diversus n. sp.

(Plates LX, LXI. LXII, LXIV)

original description

Size. Length of body from tip of head to apex of tegmen, 3.8 mm.
to 4 mm. Length of tegmen, 3.2 mm. to 3.3 mm.; width of tegmen,
2 mm.

This is a small and elongate species.

Color. Uniform, amber yellow or tan. Vertex, pronotum and
mesonotum light yellow at margins, deeper through middle. Eyes
brown. Frons uniform yellow with extreme margins dark brown.
Clypeus light yellow with somewhat darker oblique bands on each
side. Thorax and abdomen yellow tan, except basal abdominal
segments of male pale whitish yellow. Legs yellowish, tarsus
washed in reddish-brown, tips of claws black. Tegmina amber
yellow, semitranslucent, veins thick and light cream, a few dark
cross veins present in cell Cu^b-

Structural details. Vertex narrow, only twice wider than width of
eye, only slightly produced beyond eyes, its lateral margins diverg-
ing posteriorly. Frons one fifth longer than its width at posterior
margin, its lateral margins greatly elevated and somewhat diverg-
ing near anterior or apical end; a median carina present for two
thirds of its length. Length of clypeus equal to wudth of posterior

Doering: The Subfamily Issinae 533

or basal margin of the frons. Pronotum twice as long at middle as
the vertex, narrowed behind eye to a slender arm, forming a small
tuberculate knob just back of the posterior lateral angle of the eye,
but not deeply notched as in some species. Mesonotum on median
line twice as long as pronotum, a median carina faintly indicated
and two lateral ones even more faint than the middle ones. Tegmina
somewhat elongate, flat, not much inflated; costal margin more
parallel to margins of corium than in related species, veins thick
and elevated, with longitudinal veins less pronounced; clavus long,
at least twice and sometimes more than twice the distance from its
apex to apex of wings; vein Sc^ outwardly curving or at least sinuate
so that some of the costal areolets are longer than wide or at most
square; vein SCo usually two-branched, vein R frequently four-
branched, vein M four- or five-branched, all connected by cross veins
of the same length which makes it difficult to determine on which
vein the branches belong; vein Cuj branched at apex of basal third,
then the branches combining or running very close together very
near apex. Hind wings reduced to minute scales.

Male genitalia. Anal flap (10th abdominal segment) short, nar-
rowed slightly at base, then broadening posteriorly. The ventral
posterior edge of the tube extended into a rounded, lobelike flap
which is shorter than in many species. The eleventh segment show-
ing slightly beyond the dorsal edge of the tubular part as a small
ringlike segment and with a short fiingerlike stylus.

Each harpago from a flattened lateral view is an elongate lobe
with its ventral margin strongly outwardly rounded and its dorsal
margin shallowly concave through middle, after which the posterior
third of this margin is reflexed, inwardly forming a narrow pro-
jection, the caudal angle of which is bifurcate, with a dorsal process
which is bluntly rounded at end and projects inward and the ventral
process which is sharply pointed and projected ventrad and ex-
ternally.

The aedeagus is shorter than in some species, being a thick,
tubular semisclerotized process which ends bluntly at apex in two
lobes. On the left side it bears a sharply pointed, sclerotized process
attached at middle and extending to about the apex of the middle
third of the aedeagus. Just dorsad of this middle hook is a more
lobate process. A second long, curved, well-sclerotized hook, par-
tially covered by the theca, is attached somewhat near the base of
the aedeagus and extends to the same distance as the median one.
At a point where it emerges from the theca it bears a short, dorsal,
sharply pointed spine. On the right side the aedeagus shows only

534 The University Science Bulletin

a median, slightly curved process. The theca is a cylinder around
the basal third of the aedeagus, after which dorsad it extends over
the aedeagus as a lobelike hood, which is slightly broader and longer
on the left side than on the right.

Comparative notes. N. diversus is closely allied with N. uniformus
and curvaminis. The three species are difficult to distinguish ex-
ternally and the genitalia are more similar than in other species.
The latter, however, do show variations, which are constant. Many
slides of uniformus were made to prove this. The following external
differences are helpful in distinguishing the three: curvaminis is
more readily separated from the other two because the vein Sc^ is
distinctly arched inwardly, making the small cells in the costal area
wider than long, in uniformus vein Sc^ is straight and the costal cells
are square, in diversus Sc^ tends to curve outwardly, thus making
the costal cells usually longer than wide ; the number of cross veins
in the costal area tends to vary in the three species, numbering
four or five in curvaminis and diversus, and seven or eight in
uniformus; the clavus in diversus is relatively longer, being more
than twice as long as the distance from its apex to apex of tegmen,
while in curvaminis and uniformus it is only approximately twice
longer; the longitudinal veins branch more in curvaminis than in
the other two, especially M, which is five-branched in curvaminis
and only four in uniformus and diversus; in diversus the clypeus is
relatively shorter, being equal only to the basal or posterior margin
of the frons and in the other two it is equal to one lateral margin
of the frons.

Location of types. Holotype male and allotype female, collected
in the Santa Rita mountains, Arizona, by Paul Oman, June 27, 1933.
Also twenty-nine female and twenty-one male paratypes collected
during June at the same place and by the same collector. Also one
male paratype collected June 12, 1933, and two others, August 18,
1935, by R. H. Beamer. The types and most of the paratypes in
the National Museum, Washington, D. C. ; the other paratypes in
the Snow Entomological Collection, University of Kansas.

The Genus Misodema Melichar, 1906

Melichar, Leopold. Monographie der Issiden (Homoptera). Abk. k. k. Zool.-Bot. Ges.
Wien., Ill, pt. 4, 1906.

Comparative notes. The most distinctive characteristics of this
genus are the following: opaque tegmina with a coarse reticulation
of thick, strongly elevated veins, a very short clavus, as compared
to length of the tegmen and costal margin not inflected; scutellum

Doering: The Slbfamily Issinae 535

long with three prominent longitudinal carinae; wings lacking;
hind tibiae with four strong spines.

Misodema reticulata Melichar, 1906

(Plates LXVI. LXVII)

Melichar, Leopold. Monographie der Issiden (Hoinoptera). Abli. k. k. Zoijl. Bot. Ges.
Wien, III, pt. 4, 1906.

&ize. Length of body from tip of head to apex of tegmen, 3.5 mm.
to 4 mm.

Color. Uniform testaceous brown or gray-brown; tegmina same,
but approaching amber when light is transmitted through them.
Vertex light brown except anterior margin, lateral margins and
median carina, which are light yellow. Frons uniform brown with
a yellow median carina, narrow yellow lateral margins and a longi-
tudinal row of yellow spots just mesad of each lateral margin.
Pronotura brown with a cluster of round yellow spots on each
lateral half. Mesonotum dark brown at base, lighter brown areas
across apex. Tegmina opaque, uniformly dark brown, with elevated
prominent veins, which are more yellowish than the cells. Underside
of body and legs brown.

Structural characteristics. Head narrower than pronotum. Vertex
subequal in length and width, depressed through middle, a weak
median carina present, lateral margins elevated, anterior margins
triangularly produced cephalad for a considerable distance beyond
eyes, so that vertex and frons as viewed from the side form an acute
angle. Frons about one fifth longer than wide, lateral margins
rounding, greatest width across anterior third, anterior margin deeply
notched to receive postclypeus, posterior margin as viewed from the
cephalic aspect somewhat rounding. Clypeus about one half the
length of the frons, slightly inflated. Pronotum collarlike, strongly
produced anteriorly into the head, very narrowed at sides, deeply
emarginate behind, and length through middle subequal to length
of vertex at middle. Scutellum large, anteriorly greatly produced
into pronotum, narrowly tapered at sides and moderately produced
caudad to a rounded apex; three longitudinal carinae present.
Tegmina longer than body, considerably inflated, at base broadly
expanded, gradually narrowing toward apex; traversed by con-
spicuous thick veins making an irregular network so that the course
of the longitudinal veins is obscured; clavus short, with its tip
scarcely reaching to middle of anal margins, costal margin not in-
flected at base. Hind wings lacking. Hind tibiae with four spines.

536 The University Science Bulletin

Comparative notes. The outstanding structural characteristics of
the species are the heavy reticulation of the tegmina, the short
clavus, the strongly produced vertex, the collarlike pronotum, the
tricarinate scutellum and the quardrispinose hind tibiae. It re-
sembles most closely Dictyonissus griphus Uhler, but is easily dis-
tinguished from the latter by the appearance of the tegmina, which
are opaque and hairless in Misodema, but translucent and covered
with conspicuous hairs in Dictyonissus.

Location of type. National Museum, Washington, D. C.

Notes on distribution. Melichar listed this species from Mexico
and Texas. The label on the type specimen states that it was col-
lected in Texas, by C. V. Riley.

The Genus Dictyssonia, 1936

Ball, E. D'. Some New Issids With Notes on Others (Homoptera, Fulgoridae). Proc.
Biol. Soc. Wash. 49: 155-158, 19.S6.

Comparative notes. Doctor Ball in his description of the genus
states that this genus is the size and form of Dictyobia Uhler and
resembles it in several respects. It is separated from this genus by
having definite bullae at the outer angles of the tegmina, uniformly
finely reticulate tegmina, in which the major venation is almost
lost in the reticulation and the posterior margins rounding together.
The genus is separated from Neaethus Stal, Doctor Ball points out,
by having tegmina much longer and narrower behind and finer
reticulation and from the genus Dictyssa likewise, by having this
finer reticulation.

In wing venation this genus fits in the tribe Issini rather than
Hysteropterini, in which it is placed in the present key mainly be-
cause it has only vestiges of hind wings. The wing venation is as
follows: Scj lacking apparently, altho the veins forming the dark
reticulation run together at the place where Sc^ would be found and
simulate a longitudinal vein; veins SCo and R are fused for a short
distance after leaving the main vein trunk and vein Sc, appears
to have two branches and R three; vein M appears to have six
branches, but with all of these three veins the longitudinal branches
are not easily distinguished in the apical reticulation. Vein Cu^ is
distinctly a single vein and in this respect resembles the Issini and
Thioniini. The hind tibiae bear two lateral spines.

There is only one species in the genus.

Doering: The Subfamily Issinae 537

Dictyssonia beameri Ball, 1936

(Plates I.X\T. I.XVII)

Comparative notes. This is a medium-sized Issid, measuring
4.8 mm. to 5 mm. in length, testaceous-brown in color with semi-
translucent, whitish-gray tegmina, thickly traversed with reddish-
brown longitudinal veins and a fine pitch brown reticulation, so that
superficially the insect has a peppered appearance.

In his description Doctor Ball states that this species is the size
of Dictijobia permutata Uhl., but is distinguished by having the
elytra more inflated and rounding together behind, with definite
bullae. It likewise has a finer reticulation, whiter wings, and much
narrower vertex than this species.

Location of types and notes on distribution. Described from three
examples taken in the Pinery Canyon, in the Chircahua mountains
in Arizona. Holotype male and one paratype male in Doctor Ball's
collection, Tucson, Ariz., and one paratype in the Snow Entomo-
logical Collection at the University of Kansas.

538 The University Science Bulletin

PLATE XLIX

1. Lateral aspect of UUxes scutatus Walker.

2. Dorsal aspect of Ulixes scutatus Walker.

3. Lateral aspect of Traxus juhnis Metcalf.

4. Dorsal aspect of Traxus fulvm Metcalf.

5. Dorsal aspect of Tylana ustulata L'hler.

6. Dorsal aspect of head and thorax of Euthiscia

tuberculata Van Diizee.

7. Lateral aspect of Euthiscia tuberculata Van Duzee.

Doering: The Subfamily Issinae

539

PLATE XLIX

I. Ulixcs sculolus

5 Tytono ustulQta

Euthiscia tuberculoto

7. Euttiiscio tuberculoto

540 The University Science Bulletin

PLATE L

1. Frontal aspect of head of Picumna ovatipennu Walker, drawn from
homotype in British Museum Collection.

2. Ventral aspect of female abdomen of Picumna ovatipennis Walker,
drawn from homotype in British Museum Collection.

3. Dorsal aspect of Picumna ovatipennis Walker, drawn from homotype
in British Museum Collection.

4. Dorsal aspect of Picumyta maculata (Melichar).

5. Dorsal a.spect of Picumna chinoi n. sp.

6. Dorsal aspects of Thionia simplex (Germar).

Doering: The Slbfamily Issinae

541

PLATE L

posienor morqin

frons

clypeus

I. Picumno ovatipennis

2. Pi cum no ovatipernis

3. Picumno ovatipennis

4 Picumno maculato

5 Picumno chinaf

e.Thionio simplex

542 The University Science Bulletin

PLATE LI

1. Dorsal aspect of Thionia oviani n. sp.

2. Same view of Thiania producta Van Duzee.

3. Same view of Thionia quinquata Metcalf.

4. Same view of Thionia bullata Say.

5. Same view of Thionia naso Fowler.

6. Same view of Thionia elliptica (Germar).

Doeking: The Subfamily Issinae

543

PLATE LI

I. Thioria omani

3. Thionia quinquo ta

2 Thionia productq

4. Thionia bjllata

5. Thionia naso

6. Thionia elliphco

544 The University Science Bulletin

PLATE LK

1. Frontal aspect of Ulixes scutatus Walker.

2. Same view of Troxus jiihnis Metcalf.

3. Same view of Tylana tistulata Uhler.

4. Same view of Picumnn chinai n. sp.

5. Same view of Thionin siyyiplex (Germar).

6. Same view of Thionia elliptica (Germar).

7. Same view of Thionin hullata Say.

8. Same view of Thionia producta Van Duzee.

9. Same view of Thionia omani n. sp.

10. Same view of Thionia na.so Fowler.

11. Same view of Euthiscia iuberculata Van Duzee.

12. Same view of Thionia quinqiiata Metcalf.

13. Same view of Piciimna macidata (Melichar).

Doering: The Subfamily Issinae

545

PLATE LII

i::^" posterior margin

frons

2 Troxus fulvus

Ulixes scutatus

l3.Picumna maculato

4 Picumna chinai

5.ThioTiiQ simolen

e.Thlonio ellipfica

8- 9.Thionia omani

Thionia oroducta

10.

Thicnia noso

"■Euthiscio tuberculata

l2 Thionio quinquato

35—4141

546 The University Science Bulletin

PLATE LIII

1. Flattened dorsal view of anal segments (10th and

11th) of Traxus fulvus Metcalf.

2. Same view of Picumna maculata (Melichar).

3. Same view of Picumna chinai n. sp.

4. Same view of Thionia naso Fowler.

5. Same view of Thionia elliptica (Germar).

6. Same view of Thionia -producta VanDuzee.

7. Same view of Thionia simplex (Germar).

8. Same view of Euthiscia tuberculata VanDuzee.

9. Same view of Tylana ustulata Uhler.
10. Same view of Thionia bullata Sav

Doering: The Subfamily Issinae

547

PLATE LIII

) ,^JOth segment

•-.'^-.;_ ^— nth segment' ,

^_ stylus ._}''
' onol flap

.Traxus fulvus

5. Thionia ellipfica

2.Picumna maculata

3.Picumna chinoi

i

4 .Ttiionia naso

T.Ttiionia simplex

e.Ttiionio product o

S.EuthisciQ tuberculato

9. Tylana ustulato

10. Thionia bullata

548 The University Science Bulletin

PLATE LiV

1. Left view of aedeagus of Picumna chinai n. sp.

2. "Ventral view of theca and aedeagus of Picumna chinai n. sp.

3. Same view of Picumna maculata (Melichar).

4. Left view of theca and aedeagus of Tylana ustulata Uhler.

5. Same view of Picumna maculata (Melichar).

6. Ventral view of theca and aedeagus of Euthiscia tuberculata VanDuzee.

7. Same view of Tylana ustulata Uhler.

8. Same view of abdomen showing aedeagus of Ulixes scutatus Walker,
drawn from type material from British Museum Collection.

9. Left view of theca and aedeagus of Euthiscia tuberculata VanDuzee.

10. Dorso-posterior view of abdomen and aedeagal structure of Ulixes
scutatus Walker from the tj-pe material in the British Museum Collection.

Doerixg: The Subfamily Issinae

549

PLATE LIV

fO.UIixes scutatus

550 The University Science Bulletin

PLATE LV

1. Left view of aedeagtis and theca of Thionia naso Fowler.

2. Same view of Thionia si^nplex (Germar).

3. Same view of Traxus fulvus Metcalf.

4. Same view of Thionia elliptica (Germar).

5. Ventral view of aedeagus and theca of Traxus fulvus Metcalf.

6. Same view of Thionia elliptica (Germar).

7. Same view of Thionia producta VanDuzee.

8. Left view of aedeagus and theca of Thionia producta VanDuzee.

9. Ventral view of aedeagus and theca of Thionia bullata Say.
10. Left view of aedeagus and theca of Thionia bidlata Say.

Doering: The Subfamily Issinae

551

PLATE LV

552 The University Science Bulletix

PLATE LVI

1. Flattened lateral view of harpago of Picumna chinai n. sp.

2. Same view of Picumna maculata (Melichar).

3. Same view of Euthiscia tuberculata VanDuzee.

4. Same view of Tylana ustulata Uhler.

5. Same view of Thionia simplex (Germar).

6. Same view of Thionia producta VanDuzee.

7. Same view of Thionia elliptica (Germar).

8. Same view of Traxus fulmis Metcalf.

9. Same view of Thionia bullata Say.
10. Same view of Thionia naso Fowler.

Doering: The Subfamily Issinae

553

PLATE LVI

554 The University Science Bulletin

PLATE LVII

1. Lateral view of Hystcroplcrum auroreum Uhler.

2. Dorsal view of Hysteropterum sepulchralis Ball.

3. Lateral view of Hysteropterum sepulchralis Ball.

4. Dorsal view of Hysteroplerum auroreum Uhler.

5. Dorsal view of Hysteropterum cornutum Melichar.

6. Lateral view of Hysteropterum cornutum Melichar.

7. Lateral view of Hysteropterum unum Ball.

S. Dorsal view of Hysteropterum jusco77iaculosum n. sp.

9. Lateral view of Hysteropterum, juscomaculosuin n. sp.

10. Dorsal view of Hysteropterum unum Ball.

11. Lateral view of Hysteropterum, puntiferuin Walker.

12. Dorsal view of Hysteropterum puntijerum Walker.

13. Lateral view of Hysteropterum bufo VanDuzee.

14. Dorsal view of Hysteropterum bufn VanDuzee.

Doeeixg: The Subfamily Issinae

555

PLATE LVn

. 2.H.sepulchralis

3.H sepulchral is

6 . Hvsterofjteram (.ornulum

10 Hunum

l2.H,punctifetum

I3.H. bufo

14. H bufo

556 The University Science Bulletin

PLATE LVIII

1. Frontal aspect of head of Hysteropteru77i unum Ball.

2. Same aspect of Hysteropterum sepulchralis Ball.

3. Same aspect of Hysteroterum bufo VanDuzee.

4. Same aspect of Hysteropterum puntijerum Walker.

5. Same aspect of Hysteropterum auroreum Uhler.

6. Same aspect of Hysteropterum fuscomaculosum n. sp.

7. Left view of aedeagus and theca of Hysteropterum

puntijerum Walker.

8. Frontal aspect of head of Hysteropterum cornutum

Melichar.

9. Left view of aedeagus and theca of Hysteropterum

fuscomaculosum n. sp.

10. Same view of Hysteropterum sepulchralis Ball.

11. Same view of Hysteropterum cornutum, Melichar.

12. Same view of Hysteropterum auroreum Uhler.

13. Same view of Hysteropterum unum Ball.

Doering: The Subfamily Issinae

557

PLATE LYIII

558

The University Science Bulletin

PLATE LIX

1. Flattened dorsal view of anal segments (10th and 11th)

of female of Hysteropterum unuvi Ball.

2. Same view of male of Hysteropterum unum Ball.

3. Same view of female of Hysteropterum auroreum Uhler.

4. Same view of male of Hysteropterum auroreum Uhler.

5. Same view of male of Hysteropterum cornutum Melichar.

6. Same view of female of Hysteropterum cornutum Melichar.

7. Same x'lew of female of Hysteropterum, fuscomaculosum n. sp.

8. Same view of male of Hysteropterum jnintijerum Walker.

9. Same view of male of Hysteropterum sepulchralis Ball.

10. Same view of female of Hysteropterum sepulchralis Ball.

11. Same ^■iew of male of Hysteropterum, juscom.aculosum n. sp.

12. Same view of female of Hysteropterum puntiferum. Walker.

13. Flattened lateral view of harpago of Hysteropterum

unum, Ball.

14. Same view of Hysteropterum sepulchralis Ball.

15. Same view of Hysteropterum cornutum Melichar.

16. Same view of Hysteropterum punctijerum Walker.

17. Same view of Hysteropterum fuscomaculosum n. sp.

18. Same view of Hysteropterum auroreum Uhler.

Doerixg: The Subfamily Issinae

559

PLATE LIX

I. H. unum

5 . H cornu turn

9.Hsepu Ichralis

2.H.unum o

g.H.cornututi

lO.Hsepulchralis

3. H.ouroreum

7. H f uscomaculosum

^, H.auroreu

jS.H.punctiferum

I.H. foscomaculosum /"W^ i ?

12. H.punctiterum

13 H. unum

14. H.sepulchralis

15. H.cornu lum

16. Hpunctif erum

17. Htuscomaculosus

560 The University Science Bulletin

PLATE LX

1. Lateral aspect of Neaethus jenestratus Melichar.

2. Same view of Neaethus curvaminis n. sp.

3. Same view of Neaethus diversns n. sp.

4. Same view of Neaethus uniformus n. sp.

5. Same view of Neaethus similis n. sp.

6. Same view of Neaethus perhicidus n. sp.

7. Same view of Neaethus fragosus VanDuzee.

8. Same view of Neaethus sinehamatus n. sp.

9. Same view of Neaethus maculatus Melichar.

10. Same view of Neaethus intri-pennis Stal.

11. Same view of Neaethus nigronervosus Melichar.

12. Same view of Neaethus grossiis Melichar.

13. Same view of Neaethus jacintiensus n. sp.

Doering: The SrBFAMiLY Issinae

561

PLATE LX

l2N.Qro5sus

36—4141

562 The University Science Bulletin

PLATE LXI

1. Dorsal aspect of head and pronotuni of Xcaethus

unijorinus n. sp.

2. Same view of Neaethus curvaminis n. sp.

3. Same view of Neaethus diversus n. sp.

4. Same view of Neaethus jenestratus Melichar.

5. Same view of Neaethus nigronervosus Melichar.

6. Same view of Neaethus siTnilis n. sp.

7. Same view of N^caethus maculatus MeHchar.

8. Same view of Neaethus jacintiensus n. sp.

9. Same ^•iew of Neaethus perlucidus n. sp.

10. Same view of Neaethus vitripennis Stal.

11. Same view of Neaethus sinehamatus n. sp.

12. Same view of Neaethus grossus Melichar.

13. Frontal aspect of head of Neaethus uuiformus n. sp.

14. Same view of Neaethus diversus n. sp.

15. Same view of Neaethus Jenestratus Melichar.

16. Dorsal aspect of head and pronotum of Neaethus

fragosus VanDuzee.

17. Frontal aspect of head of Neaethus nigronervosus

Melichar.

18. Same view of Neaethus curvaminis n. sp.

19. Same view of Neaethus sinehamatus n. sp.

20. Same view of Neaethus gr-ossus Melichar.

21. Same view of Neaethus similis n. sp.

22. Same view of Neaethus maculatus Melichar.

23. Same view of Neaethus perlucidus n. sp.

24. Same view of Neaethus vitripennis Stal.

25. Same view of Neaethus jacintiensus n. sp.

26. Same view of Neaethus fragosus VanDuzee.

Doering: The Subfamily Issinae

563

PLATE LXI

l.N.uniformus

2 .N cur vaminis 3.N.diversus 4.N. fenestratus

5.N. nigronervosus

6 . N . s i m 1 1 i s

9.N.perlucidus lO.N. vif ripennis 1 1 .N.sinehamatus l2.N.grossus

l3.N.un formus

14.N.diversus 15. N. fenestratus

16. N. fragosus

le.N.curvaminis 19. N.sinetiamatus

N. nigronervosus

20.N.grossus

22 V N maculatus
SS.N.jacint iensus

23. N perlucidus 24.' N.vitripennis

26.' N. fragosus

564 The University Science Bulletin

PLATE LXII

1. Flattened dorsal view of anal segments (10th and

11th) of male of Neaethus unifonnu-s n. sp.

2. Same view of Ah'aethus diversus n. sp.

3. Same view of Neaethus fenestratus Melichar.

4. Same view of Neaethus curvaminis n. sp.

5. Same view of Near thus grossus Melichar.

6. Same view of Neaethus similis n. sp.

7. Same view of Neaethus vitripcnnis Stal.

8. Same view oi Neaethus fragosus VanDuzee.

9. Same view of Ncncllnis jacintiensus n. sp.

10. Same view of Neaethus perlucidus n. sp.

11. Same view of Neaethus nigronervosus Melichar.

12. Same view of Neaethus slnchamatus n. sp.

13. Flattened lateral \ipw of harpago of Neaethus

nigronervosus Melichar.

14. Same view of Neaethus maculatus Melichar.

15. Flattened dorsal view of anal segment of male of

Neaethus tnaciilatus Melichar.

16. Flattened lateral \ iow of harpago of Neaethus

similis n. sp.

17. Same view of Ah^aethus uuijunnus n. sp.

18. Same view of Neaethus diversus n. sp.

19. Same view of Neaethus curvaminis n. sp.

20. Same view of A'Caethus fe)testratus Melichar.

21. Same view of Neaethus fragosus VanDuzee.

22. Same view of Neaethus grossus Melichar.

23. Same view of Neaethus jacintiensus n. sp.

Doering: The Subfamily Issinae

565

PLATE LXII

21.

N frogosus

566 The University Science Bulletin

PLATE LXIII

1. Left view of aedeagus and theca of Ncaelhus pcrlucidus n. sp.

2. Right view of aedeagus and theca of Neaethus perlucidus n. sp.

3. Left view of aedeagus and theca of Neaethus sinehamatus n. sp.

4. Right view of aedeagus and theca of Neaethus vitripennis Stal.

5. Right view of aedeagus and theca of Neaethus sinehamatus n. sp.

6. Left view of aedeagus and theca of Neaethus vitripe^mis Stal.

7. Flattened lateral view of harpago of Neaethus sinehamatus n. sp.

8. Same view of Neaethus perlucidus n. sp.

9. Same view of Neaethus litripennis Stal.

Doering: The Subfamily Issinae

567

PLATE LXIII

508 The Uxiversity Science Bulletin

PLATE LXIV

1. Left view of aedeagus and theca of Neaeth2is curvaminis n. sp.

2. Right view of aedeagus and theca of Neaethus curvaminis n. sp.

3. Left view of aedeagus and theca of Neaethus diversus n. sp.

4. Right view of aedeagus and theca of Neaethus diversus n. sp.

5. Left view of aedeagus and theca of Neaethus iiniformiis n. sp.

6. Right view of aedeagus and theca of Neaethus uniformus n. sp.

7. Left view of aedeagus and theca of Neaethus fenestratus Melichar.

8. Right view of aedeagus and theca of Neaethus fenestratus Melichar.

9. Left view of aedeagus and theca of Neaethus jragosus VanDuzee.
10. Right view of aedeagus and theca of Neaethus fragosiis VanDuzee.

Doerixg: The Subfamily Issinae

569

PLATE LXIV

S.N.curvominis

3.N diversus

4. N diversus

6.N.uniformus

^^:^

8. N.feneslrotus

570 The University Science Bulletin

PLATE LXV

1. Left view of aedeagus and theca of Neaethus simiUs n. .sp.

2. Right view of aedeagus and theca of NeaethuH shnilis n. sp.

3. Left view of aedeagus and theca of Neaethus nigronervosus

Melichar.

4. Right view of aedeagus and theca of N^eaethns ')ugr(meri'os}is

Mehchar.

5. Left view of aedeagus and theca of Neaethus maculatus Melichar.

6. Right view of aedeagus and theca of Neaethus maculatus

Melichar.

7. Left view of aedeagus and theca of Neaethus grossus Melichar.

8. Left view of aedeagus and theca of Neaethus jacintiensus n. sji.

9. Right A'iew of aedeagus and theca of Neaethus grossus Melichar.
10. Right view of aedeagus and theca of N^eacthus jncivtien-vis n. sp.

Doering: The Subfamily Issinae

571

PLATE LXV

I.N^Imilis

Z N.similis

S.Nnijronervosus

4.N.niqroneryosus

572 The University Science Bulletin

PLATE LXVI

1. Right view of theca and aedeagus of Dictyunia obscura Uhler.

2. Right view of thcca and aedeagus of Dictyonissus griphus Uhler.

3. Left view of theca and aedeagus of Dictyonissus griphus Uhler.

4. Left view of theca and aedeagus of Diclyonia obscura Uhler.

5. Right \iew of thcca and aedeagus of Dictyonissiis nigropilosus n. sp.

6. Left view of theca and aedeagus of Dictyonissus nigropilosus n. sp.

7. Lateral aspect of Dictyonia obscura Uhler.

8. Lateral aspect of Misodcma reticulata Melichar.

9. Dorsal aspect of Misodema reticulata Melichar.
10. Lateral aspect of Dictyonissus griphus Uhler.
IL Dorsal aspect of Dicfyoni,ssHS griphus Uhler.

12. Dorsal aspect of Dictyonissus nigropilosus n. sp.

13. Dorsal aspect of Dictyssonia beameri Ball.

Doering: The Subfamily Issinae

573

PLATE LXVI

II. Dictyonissus grlphus

)2. Dictyonissus nigropilosus

13 D'ctyssonia beomeri

574 The University Science Bulletin

PLATE LXVII

1. Frontal as])ect of head of Dictijonissiis giiphus Uhler.

2. Same view of Dictyonissus nigropilosus n. sp.

3. Same view of Misodema reticulata Melichar.

4. Same view of Dictyssonia beameri Ball.

5. Flattened lateral view of harpago of Dictyonissus nigropilosus n. sp.

6. Dorsal view of head and pronotum of Dictyonia obscura Uhler.

7. Frontal aspect of head of Dictyonia obscura Uhler.

8. Flattened lateral view of harpago of Dictyonia obscura Uhler.

9. Same view of Dictyonissus griphus Uhler.

10. Dorsal view of anal segments of male and female of Dictyonia obscura

Uhler.

11. Ventral view of abdomen of male of Dictyonissus griphus Uhler.

12. Dorsal view of anal segments of male of Dictyonissus griphus Uhler.

13. Ventral view of abdomen of male of Dictyonia obscura Uhler.

14. Ventral view of abdomen of female of Dictyonissus griphus Uhler.

15. Dorsal view of anal segments of male of Dictyonissus nigropilosus n. sp.

16. Ventral view of abdomen of female of Dictyonia obscura Uhler.

Doerixg: The Subfamily Issinae

OID

PLATE LXVII

I. 2.Dict yonissus Misodema reticulata

Dictyonissus griphus nigrooilosus

5. D-nigropilosus

II. D. griphus

4 D'ictyssonia beomeri

7 Dictyonia obscura

9 0 griphus

10.
D. obscura

12. D. griphus

13. D- obscura

4.D. griphus 15. D. nigropilosus

\6.^

obscura

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 21

The Morphology of the Carolina Mantis

PHILIP LEVEREAULT,

Department of Entomology, University of Kansas

Section II: The Musculature*

Abstract: In this section of the study of the morphology of the Carolina
mantis attention is concentrated upon the description of the musculature, but
not exclusively, for the important functions of the muscles, obvious from dis-
sections of fixed specimens, are discussed. The text is supplemented by twelve
plates of penned drawings.

CONTENTS

IXTRODUCTION

P.ART 1. The Ceph.^lic Muscles: page

Facial muscles 579

Antennal muscles 579

Stomodeal muscles 580

Mandibular muscles 581

Maxillary muscles 582

Labial muscles 583

Plates:

LXVni 587

LXIX 589

Part 2. The Thor-acic Muscles:

Neck muscles 590

Prothoracic muscles 592

Anterior somitic muscles 592

Leg muscles 594

Posterior somitic muscles 595

Pterothoracic muscles 596

Mesothoracic somitic muscles 597

Mesothoracic leg muscles 598

Metathoracic somitic muscles 600

Metathoracic leg muscles 601

* Continued from Scienre Bull. Vol. XXIV, pp. 203-259, and submitted with it to the De-
partment of Entomology and the faculty of the Graduate School of the University of Kansas
in partial fulfillment of the requirements for the degree of doctor of philosophy. Part three
to follow.

37—4141 (577)

578 The University Science Bulletin

Plates : pagh

LXX 605

LXXI 607

LXXII 609

LXXIII 611

LXXIV A 612

LXXIV B 613

LXXV 615

Part 3. The Abdominal Muscles:

Muscles of the first segment 616

Muscles of the second segment 617

Muscles of the third segment 618

Table of equivalents of the fourth, fifth and sixth segments 619

Muscles of the female terminalia 619

Muscles of the male terminalia 622

Plates :

LXXVI 627

LXXVIl 629

LXXVIII 631

LXXIX 633

INTRODUCTION

FOR the past decade the close relationship of the insect skeleton
and the musculature has been strongly emphasized. Some in-
vestigators insist that this relationship is so close the two structural
systems must be discussed together under one heading. The dis-
covery of the morphological and physiological relationship of the
skeleton and musculature, as lately interpreted, is a worthy con-
tribution to insect morphology, but it should not be run out of
bounds. Scientific concepts should not be twisted into dogmas.

The functional intimacy between the skeleton and musculature
does not force the musculature into a subservient morphological
position. The musculature is derived from mesodermic embryonic
tissue, a tissue different from the epidermis which forms the skele-
ton. In addition, the mode of development, the formation of coe-
lomic sacs and the splitting into muscle groups, is very distinctive.
Such fundamental differences warrant the consideration of the mus-
culature as a morphological system coordinate with the skeleton,
and in this study of the Carolina mantis, worthy of as detailed a
treatment.

Levereault: The Carolina Mantis 579

PART 1— THE CEPHALIC MUSCLES

The cephalic muscles of the Carolina mantis may be assigned to
three groups, the facial, stomodeal, and gnathocranial muscles. With
the exception of the antennal, all of the cephalic muscles of this in-
sect are concerned with the function of feeding, directly or in-
directly.

In the facial group there are six pairs of muscles:

1. The labral compressors (figures 1 and 8) ; a pair of V-shaped bundles ex-
tending from the basimedial region of the labrite to the adoral wall of
the labrum (see figure 8). The contraction of these muscles flattens the
basal portion of the labrum, thereby widening the prebuccal space.

2. The anterior labrals (figure 1) ; a pair of slender ribbonlike bands extend-
ing from the frontal sclerite, just below the median ocellus, to the median
portion of the basal margin of the labrite. The labrum is extended for-
ward by a short contraction of these muscles.

3. The posterior labrals (figure 1); slender, flat bands extending from the
frontal sclerite, laterad of the anterior labral muscles, to the lateral
tongues of the labrite (the so-called "tormae"). A short contraction of
these muscles draws back the labrum against the mandibles. If the an-
terior and posterior labral muscles are fully contracted together, the
labrum is retracted within the lower part of the epicranium. This re-
traction is frequently performed, especially when the mantis reaches for
the entrails through a break in the skeleton of its victim.

1. The antennal levator* (figures 1 and 2) ; a short, wedge-shaped muscle
extending from the anterior transtral arm to the upper part of the basal
rim of the first antennal segment.

5. The median antennal depressor (figures 1 and 2) ; a short muscle extend-
ing from the anterior transtral arm to a tendon near the median portion
of the basal rim of the first antennal segment.

6. The lateral antennal depressor (figures 1 and 2) ; a short muscle extend-
ing from the anterior transtral arm to the lateral portion of the basal
rim of the first antennal segment.

The mechanism of the base of the antenna of this insect is an in-
teresting one. The skeleton of the basal, or first, segment is articu-
lated with the antennifer (the toothlike projection between muscles
5 and 6, figure 2: for more skeletal details see section I, Plate IX,
figure 2), forming a single point articulation which permits consid-
erable freedom of movement. The three muscles, 4, 5, and 6, are
attached at the most advantageous points of the basal rim of the
first segment, enabling the mantis to swing the entire appendage in
a circle by variations of contractions.

* For ease in discussion it will be assumed that ail muscles are paired, unless specifically
designated as unpaired.

580 The University Science Bulletin

Within the antenna are two paired muscles:

7. The extensor of the flagcHum (figure 7) ; a twinned muscle extending
from the lateroventral basal region of the first antennal segment to the
lateral portion of the basal rim of the second, or intermediate antennal
segment. This muscle does not extend the second segment and the flagel-
lum perfectly laterally, for the basal attachment of the muscle forces the
muscle to pull the two distal segments partly downward.

8. The flexor of the flagellum (figure 7) ; a twinned muscle extending from
the mediodorsal basal region of the first segment to the median portion of
the basal rim of the second segment. The flexion of the second segment
and the flagellum, like the extension, is not perfectly lateral because the
basal attachment of the flexor forces the muscle to pull partly upward.

The buccal, pharyngeal, and esophageal portions of the insect ali-
mentary tract are relatively indefinite regions of the stomodeal in-
vagination, and muscles acting upon these regions may well be
called stomodeal muscles in the Carolina mantis. In figure 8 only
the left members of the pairs are shown.

9. The anterior buccal dilators (figure 8) ; two groups, usually four in each,
of muscle bundles extending from the median area of the subfrontal
sclerite to the anterior wall of the buccal region.

10. The first dilators of the pharynx (figure 8) ; a pair of short, flat muscles
extending from the lower medial area of the frontal sclerite to the an-
terior wall of the pharynx.

11. The second dilators of the pharynx (figure 8) ; a pair of long muscles ex-
tending from the mediodorsal area of the frontal sclerite to the anterior
pharyngeal wall.

12. The third dilators of the pharynx (figures 1 and 8) ; a pair of strong
muscles extending from the ocular ridges near the anterior transtral arms
to the sides of the pharynx.

13. The fourth dilators of the pharynx (figure 8) ; a pair of flat, fan-shaped
muscles extending from the mediodorsal area of the frontal sclerite to the
upper wall of the pharynx.

14. The fifth dilators of the pharynx (figure 8) ; a pair of long, flat, fan-
shaped muscles extending from the apex of the frontal sclerite to the
upper wall of the pharynx.

15. The first posterior buccal dilators (figure 8) ; a pair of short, thick muscles
extending from the anterior bar of the transtral body to the posterior
wall of the buccal region.

16. The second posterior buccal dilators (figure 8) ; a pair of small, thick
muscles extending from the anterior bar of the transtral body to the pos-
terior wall of the buccal region.

17. The sixth dilators of the pharynx (figure 8) ; two groups of thick, closely
set muscle bundles extending from the upper surface of the anterior bar
of the transtral body to the lower wall of the pharynx.

18. The seventh dilators of the pharynx (figure 8) ; two flaring muscles
extending from the upper surface of the posterior transtral arms to the
posterior lateral walls of the pharynx. These muscles, acting alone,
flatten the po.sterior part of the pharynx, but, since the circular muscles

Levereault: The Carolina Mantis 581

of this region are the primary constrictors, these seventh dilators are
the antagonists to the circulars. However, to perform a dilation of this
stomodeal region, these muscles must act in unison with the fifth
phaiyngeal dilators.

All of these stomodeal dilator muscles are the antagonists of the
stomodeal circulars. When food is passed up before the mandibles
the labral compressor muscles (1) flatten the labrum, enlarging the
preoral space. The buccal dilators (9, 15, 16) contract to enlarge
the mouth region for the reception of the food. After the food has
entered the mouth the dilators relax and the buccal circular muscles
contract, forcing the food backward in the stomodeum. As the
buccal circulars contract the pharyngeal dilators (first 10, 11, 12,
with 16 and 17, then 13 with 17, and finally 14 with 18) pull succes-
sively upon the pharyngeal walls. As the food is drawn posteriorly
by the action of the dilators, the pharyngeal circular muscles force
it backward. In the living animal these motions of dilation and
contraction make the stomodeum ripple as bolt after bolt of food is
forced toward the stomach.

The gnathocranial muscles are a complicated group, moving the
cephalic postoral appendages and the salivary pocket:

19. The mandibular flexor (figure 3) ; a large, complex muscle extending
from the dorsal area of the epicranium to the flexor tendon of the mem-
brane near the mesal portion of the mandibular basal rim.

20. The mandibular extensor (figure 3) ; a fiat, flaring muscle extending from
the postocular area of the genal sclerite to the extensor tendon of the
membrane near the lateral portion of the mandibular basal rim.

21. The fiexor-adjustor of the mandible (figure 6) ; a short, stout muscle, ex-
tending from the outer surface of the anterior transtral arm to the in-
flection on the posterior surface of the mandible.

The flexor-adjustor muscles aid the mantis in breaking hard-
shelled prey. They do not contribute a strong flexion for mastica-
tion, but they help to some degree. Their principle function is to
produce an anterior-posterior movement to the mandibles. There is
but a slight movement possible in this direction because of the rel-
atively closely held articulatory points, but this slight movement
is important. When the mantis tries to bite through a particularly
testy beetle it contracts the large flexor muscles (19), holds, and
then the flexor-adjustors, contract alternatingly, giving the incisor
processes a gnawing effect. These movements are quite comparable
to our own efforts in dealing with the ordinary cafe steak.

22. The vestigial ventral adductor of the mandible (figure 6) ; a flat, wedge-
shaped muscle extending from the midlateral area of the mandibular
wall to the glottal sclerite 2 (see section I, Plate X, figures 11 and 14).

582 The University Science Bulletin

The vestigial ventral adductors have been homologized by Snod-
grass with the ventral adductors of myriopods, diplopods, certain
crustaceans and apterygotans. In the Carolina mantis these muscles
have survived their usefulness. They do play a minor role, which
will be discussed at another point.

The following muscles move the maxilla and its parts:

23. The cardinal rotator (figures 9 and 11); a three-part muscle extending
from the posterior portion of the ocular ridge, 23a, and 23b, from the
lateral region of the occipital sclerite, 23c, to the tendon of the cardinal
process, 23a and 23c, and to the small tendon of the membrane near the
cardinal process, 23b.

In section I, I interpreted the action of these muscles as an ex-
tension. This interpretation is incorrect. There is a slight lateral
extension, but this movement is subordinate to the rotation of the
cardo. In the rotation of the cardo 23a and 23b pull the cardinal
process up toward the ocular ridge. As the cardinal process is pulled
dorsally, the outer rim of the cardo is turned ventrally, and 23c
completes the rotation by drawing the tendon of the cardinal process
against the occipital sclerite. This rotation of the cardo gives the
entire appendage a promotion, if the other maxillary muscles re-
main relaxed.

24. The median cardinal flexor (figure 12) ; a stout muscle extending from
the outer surface of the transtral body to the median region of the
cardo.

25. The distal cardinal flexor (figure 12) ; a two-part muscle extending
from the outer surface of the transtral body to the distal portion of the
cardinal rim and the upper lateral region of the stipes.

Both cardinal flexors use the cone (see section I, Plate X, figure
16, d) of the basal rim of the cardo as the articulatory point against
the epicranium. The cardinal rotator begins the flexing movement
by turning the cardo into an advantageous position. The rotation
and the subsequent contraction of the flexor muscles extend the
maxillary tip ventrally, and toward the median plane of the trunk.

26. The forcipital flexor (figures 9, 11, 12); a three-part muscle extending
from the lateral edge of the stipes, 26a, from the upper medial region
of the stipes, 26b, from the postocular portion of the genal sclerite, 26c,
to the tendon of the membrane above the base of the forceps; 26c aids
in flexing the maxilla as well as retracting it (see position of maxilla in
figure 9).

27. The stipital flexor (figures 12, 13) ; a large muscle extending from the
outer surface of the transtral body to the stipital ridge.

28. The accessory stipital flexor (figures 10, 13) ; a flat muscle extending
from the outer surface of the transtral body to the median area of the
stipes.

Levereault: The Carolina Mantis 583

The maxilla of this insect is not extended laterally to any great
degree. Most of the muscles are flexors, and, except for the partial
extension performed by the cardinal rotator, the return of the
maxilla to the relaxed position depends upon the resistance of the
skeleton to the flexing action. In observing the living mantis I
found that the maxilla is moved in a vertical direction, nearly
parallel with the sagittal plane of the trunk. 26c lifts the append-
age, and the combined action of cardinal rotator, cardinal flexors,
and stipital flexors, extends the appendage ventrally and slightly
toward the sagittal plane. This alternating retraction and ex-
tension, with the accompanying flexion and extension of the palp,
gives the mantis the appearance of a mumbling old lady.

29. The flexor of the maxillary palp (figiii-e 13) ; a small, flat muscle ex-
tending from the stipital ridge to the lower part of the basal rim of the
first palpal segment.

30. The reductor of the maxillary palp (figure 13) ; a small, flat muscle
extending from the stipital ridge to the posterior ])ortion of the basal
rim of the first palpal segment.

31. The extensor of the maxillary palp (figure 13); a small, flat muscle ex-
tending from near the stipital ridge to the upper portion of the ba.sal rim
of the first palpal segment.

There is no definite articulatory point between the basal palpal
segment and the stipes, but the intervening membrane functions as
an articulation. Its flexibility permits a free rotation, which is
effected by variations in intensity of contraction of the three basal
muscles.

32. The extensor of the second palpal segment (figure 13) ; a small muscle
extending from the upper basal area of the first palpal segment to the
uiiper portion of the basal rim of the second segment.

33. The flexor of the second palpal segment (figure 13) ; a .slender muscle
extending from the lower basal region of the first segment to the lower
portion of the basal rim of the second segment.

34. 35, 36. The flexors of the third, fourth, and fifth palpal segments (figure

13) ; slender muscles extending from the lower portions of the basal rims
of these palpal segments.

37. The pollicial productor (figure 10) ; a long, flat muscle extending from
the upper portion of the stipes to the posterior basal rim of the pollex.

The labium and its parts, and the salivary pocket are moved by
the following muscles:

38. The pollicial productor (figure 14) ; a long, .slender mu.scle extending from
the lower surface of the posterior bar of the transtral body to the mem-
Ijranc above the pollicial base.

39. The mental productor (figure 14) ; a large, flat muscle extending from
the lower surface of the posterior bar of the transtral body to the

584 The University Science Bulletin

laterodistal edges of the mental plate. This muscle is the chief productor
of the labium. The action of the poUicial productor is probably negli-
gible.

40. The apical dilators of the salivary pocket (figures 5 and 14) ; a pair of
flat muscles extending from the lateral edges of the labial stipites to the
keel of the salivary pocket.

41. The basal dilators of the salivary pocket (figures 5 and 14) ; a pair of
short, flat muscles extending from the central area of the labial stipites
to the lateral edges of the salivary pocket.

42. The anterior dilators of the salivary pocket (figures 4 and S) ; a V-shaped
pair of muscles extending from the lateral glottal sclerites (section I,
Plate X, figures 14, 2) to the median line of the anterior wall of the
salivary pocket.

43. The retractors of the salivary pocket (figures 4, 5, 8) ; a pair of long, flat
muscles extending from the lower surface of the posterior bar of the
transtral body to the lateral edges of the salivary pocket.

These muscles acting upon the salivary pocket have no firm basal
points of attachment, the so-called points of "origin." When the
anterior dilators contract they pull back the lateral glottal sclerites,
though these sclerites are held forward to some degree by the ves-
tigial adductors of the mandibles. When the posterior dilators con-
tract the labium is drawn forward to some degree.

44. The stipital reductors of the labium (figure 15) ; a pair of short, flat
muscles extending from the central part of the mentum to the median
part of the basal stipital rim.

45. The extensor of the labial palp (figure 15) ; a flat, flaring muscle extend-
ing from the lateral basal corner of the stipes to the upper portion of
the basal rim of the first palpal segment.

46. The flexor of the labial palp (figure 15) ; a flat, fan-shaped group of
bundles extending from the median margin of the stipes to the lower
portion of the basal rim of the first palpal segment.

47. The extensor of the second palpal segment (figure 15) ; a group of small
bundles extending from the upper basal region of the first segment to the
upper portion of the basal rim of the second segment.

48. The flexor of the third palpal segment (figure 15) ; a slender muscle ex-
tending from the upper distal area of the first segment to the median
portion of the basal rim of the third segment.

49. The pollicial reductors (figure 15) ; a pair of slender, flat muscles extend-
ing from the central part of the stipites to the posterior portions of the
basal rims of the pollices.

50. The forcipital reductors (figure 15); a pair of long, slender muscles ex-
tending from the central part of the stipites to the posterior portions of
the basal rims of the forcipes.

Levereault: The Carolina Mantis 585

REFERENCES

There is little need for relisting the more important studies of the cephalic
musculature of insects which are to be found in Snodgrass' "Morphology and
Evolution of the Insect Head and its Appendages," Smiths. Misc. Coll., vol.
81, No. 3, 1928, and the shorter "Evolution of the Insect Head and the Organs
of Feeding," Smiths. Rept. 1931, pp. 443-489. Little of any value has appeared
since Snodgrass wrote those papers.

586 The University Science Bulletin

PLATE LXVni

Fig. 1. Facial muscles of male head.

Fig. 2. Basal muscles of right antenna.

Fig. 3. Anterior view of flexor and extensor muscles of right mandible ( 5 ).

Fig. 4. Anterior (%) view of salivary pocket with posterior wall of glotts
depending.

Fig. 5. Posterior view of salivary pocket.

Fig. 6. Basal details of musculature of right mandible.

Fig. 7. Intrinsic muscles of right antenna.

Fig. 8. Stomodeal muscles from median plane.

Levereault: The Carolina Mantis

587

PLATE LXVIII

Fig. I

Fig. 2

Fig. 6

588 The University Science Bulletin

PLATE LXIX

Fig. 9. Anterior view of muscles and attachment of right maxilla to epi-
cranium. Retracted position.

Fig. 10. Outer muscles of right maxilla.

Fig. 11. Detailed view of intracranial muscles of right maxilla.

Fig. 12. Inner muscles of right maxilla.

Fig. 13. Intermediate and palpal muscles of right maxilla.

Fig. 14. Inner muscles of labium.

Fig. 15. Outer muscles of labium.

Levereault: The Carolina Mantik

589

Fig. 14

Fig. 15

590 The University Science Bulletin

PART 2— THE THORACIC MUSCULATURE

Under this heading are included the neck muscles as well as the
muscles of the three thoracic somites. This inclusion is admittedly
an arbitrary one, for no study of adult musculature can indicate
which muscles or parts of muscles belong to the head, and which to
the prothorax.

The neck of the Carolina mantis is a very flexible body region
which permits the insect to turn its head into surprisingly many
positions. The skeleton of the neck has two pairs of articulatory
points, the anterior edges of the anterior lateral cervical sclerites
(see section I, Plate XI, figures 7 and 10), and the upper points of
the posterior lateral cervical sclerites. The anterior articulatory
points press upon the condyles of the postoccipital sclerite (poc c,
figure 1), and the posterior points upon the anterior margins of the
prothoracic prepleurites. In addition to these four points of move-
ment, two lines of flexure, which crease the posterior lateral cervical
sclerites, form angles which extend and retract the skeleton. These
features, with the cervical musculature, enable the mantis to twist
the posterior rim of the cranium by quarter turns, or to shorten or
lengthen one side of the neck.

The muscles involved in these movements are:

1. The lateral dorsal rotator of the head (figure 3) ; a three-part muscle ex-
tending from the central part of the anterior lateral cervical sclerite, la,
from the mid-upper edge of the same sclerite, lb, from the lower edge
of the anterior rim of the prothoracic tergum, Ic, to the membrane near
the dorsolateral portion of the postoccipital sclerite.

2. The protractor of the posterior lateral cervical sclerite (figures 2 and 3) ;
a strong muscle extending from the anteromesal region of the prothoracic
tergum to the lateroposterior portion of the posterior lateral cervical
sclerite. This muscle pulls the cervical sclerite forward, enlarging the
angle of the line of flexure and. thereby, elongating the neck.

3. The retractor of the anterior lateral cervical sclerite (figures 2 and 3) ; a
large, flat muscle extending from the anteromesal area of the prothoracic
tergum to the center of the anterior lateral cervical sclerite. This muscle
is the antagonist of 2. When it contracts it closes the angle of the flexure
of the posterior lateral cervical sclerite, thereby shortening the neck.
However, if the protractor, 2, contracts first, the retractor merely com-
pletes the opening of the angle of the line of flexure.

4. The lateral rotator of the head (figure 2) ; a long, slender muscle extend-
ing from the tendon of the tergal adductor of the prothoracic coxa to
the upper anterior edge of the anterior lateral cervical sclerite. This
muscle might close the angle of the posterior lateral cervical sclerite, and
turn the head laterally, but its slenderness indicates it plays a very minor
role in the movements of the neck.

Levereault: The Carolina Mantis 591

5. The median dorsal rotator of the head (figures 1 and 2) ; a slender muscle
extending from the long tendon of the membrane near the pretergite of
the meso thoracic tergum to the tendon of the dorsal cervical sclerite.

6. The lateral ventral rotator of the head (figure 2) ; a flat muscle extend-
ing from the posterior dorsolateral region of the neck skeleton to the
membrane near the postoccipital condyle. This muscle cannot perform
a ventral rotation of the head by itself, for its basal point is not even
sclerotized. However, if the median dorsal rotator, 5, pulls upon the
tendon of the dorsal cervical sclerite, yet permits itself to be stretched
as the head is turned ventrally, the dorsal wall of the neck is held taut
and the lateral ventral rotator is enabled to effect a pull.

7. The lateral dorsal rotator of the head (figure 1) ; a two-part muscle ex-
tending from the tendon of the membrane near the pretergite of the
mesothoracic tergum, 7a, from the anteromesal region of the prothoracic
tergum 7b, to the membrane adjacent to the dorsolateral portion of the
postoccipital sclerite.

8. The ventral rotator of the head (figure 1) ; a long, strong muscle extend-
ing from the tendon of the prothoracic f ureal apophyses to the ventral
process of the postoccipital sclerite.

The above interpretations of the neck muscles are interpretations
of the primary individual functions. No doubt these neck muscles
do not function to any great extent as individuals, but as members
of groups. There are five functional groups in the neck of the
Carolina mantis, protractors, retractors, lateral rotators, dorsal and
ventral rotators.

In the protractor group both protractors of the posterior lateral
cervical sclerites pull forward the lower part of the neck. This
pull widens the angles of the lines of flexure, creasing the posterior
lateral cervical sclerites, and when these angles are obtuse the
muscles 3 complete the protraction.

The retraction of the neck is effected by the combined pull of
muscles 7 and 8, or, with 4 and 5 aiding the pair 7. The retractors,
3, of the anterior lateral cervical sclerites start the shortening of
the neck by closing the angles of the lines of flexure of the posterior
lateral cervical sclerites.

A lateral rotation of the head to the right may be started by a
contraction of 3, which would fold the right posterior lateral cervical
sclerite; then, 7 and 8, or 5 or 6, or both groups together, would turn
the head to the right. Similar actions of the left-side members of
these pairs of muscles would turn the liead to the left.

A dorsal rotation of the head, from any angle between the neck
sclerites, may be produced by the contraction of the pair 7, or with
the aid of the pairs 5 and 1. The opposite movement, the ventral

592 The University Science Bulletin

rotation of the head, may be produced by the contraction of the
pair 8, or with the aid of the pair 6.

These group functions are not performed with "smart" military
precision, as the above descriptions might imply. The Carolina
mantis is a leisurely sort of insect. It may be an awkward one, but
it does try to cover its ungainliness with a quiet, slow dignity.
When it turns its head it usually does so slowly, as the functions
of the muscles blend in protraction, rotation, and retraction by
varying intensities of contraction.

The Prothok4cic Muscles

In addition to the neck muscles lying within the prothorax

proper, there are three principal groups of muscles which act upon

the tergum, pleura, and sternum of this somite. Two groups are in

the supracoxal region, and one group in the posterior region. Most

of the supracoxal muscles are basal leg muscles, for only three pairs

of small muscles act upon the tergum and pleura.

9. The tergal adductor of the coxa (figures 4 and 5) ; a large three-part
muscle extending from the mesal region of the tergum. anterior to the
supracoxal inflection, 9a, posterior to the inflection, 9b, and 9c, to the
tendon of the jileurellite.

In Dissosteira Carolina Snodgrass found the homologue of this
mantid muscle to be a single band with a promotion function. In the
mantis only 9a can impart any promotion, for the other two mem-
bers are attached too far back on the tergum, and it is very probable
9a merely aids in the adduction of the coxa.

This adduction is one of the important movements in feeding.
If the mantis catches an insect in its right foreleg, the predator
turns the coxal base not quite a quarter-turn clockwise; then, it
brings the coxa forward so that the distal portion points ahead and
away to the left of its body. The combination of this rotation and
promotion places the tibiofemoral joint, the point near which the
victim is usually held, into a convenient position for the jaws of
the mantis. The turning is performed by the muscles 12, 13, 14, and
17; the promotion is effected by 19a and 19b.

10. The tergal remotor of the coxa (figures 4, 5, and 9) ; a big two-part muscle
extending from the midlateral region of the tergum to the tendon of the
membrane near the posterior lateral portion of the basal coxal rim.

11. The anterior sternal rotator of the coxa (figures 5, 6, 7) ; a small, flat
muscle extending from the median process of the furcasternite to the
anterior portion of the basal coxal rim.

Levereault: The Carolina Mantis 593

12. The posterior sternal rotator of the coxa (figures 5 and 9) ; a short, stout
muscle extending from the outer basal surface of the furcal apophyses to
the tendon of the membrane near the lateral portion of the basal coxal
I'im.

13. The accessory tergal remotor of the coxa (figure 5) ; a long, flaring, flat
band extending from the midlateral region of the tergum to the tendon
of the membrane near the lateral portion of the basal coxal rim.

This muscle probably corresponds to Snodgrass' "67 — the second
posterior rotator of the coxa" of Dissosteira. In the elongation of
the Carolina mantis prothorax this muscle lost its basal attachment
on the spinal process of the spinasternite, and became attached to
the tergum.

14. The posterior abductor of the coxa (figm-e 5) ; a short, flat muscle ex-
tending from the lateral region of the tergum behind the supracoxal
inflection to the tendon of the membrane near the lateral portion of the
basal coxal rim.

15. The intermediate abductor of the coxa (figures 5 and 9) ; a short, flat,
fan-shaped muscle extending from the lateral region of the tergum be-
hind the supracoxal inflection to the peak of the lateral portion of the
basal coxal rim.

16. The anterior abductor of the coxa (figure 5) ; a short, slender band ex-
tending from the lateral region of the tergum behind the supracoxal in-
flection to the peak of the lateral portion of the basal coxal rim.

17. The abductor-rotator of the coxa (figures 4 and 5) ; a long, flaring muscle
extending from the upper lateral region of the tergum to the peak of
the lateral portion of the basal coxal rim. The action of this muscle
is a combined posterior rotation and abduction, with the rotation prob-
ably stronger than the abduction.

These muscles, 14, 15, 16, and 17, belong to the "M group" in
Snodgrass' generalized plan of the basal coxal musculature.

18. The sternal adductor of the coxa (figure 6) ; a wide, flat band extending
from the lower surface of the furcal ajwphysis to the mesal j^ortion of
the basal coxal rim.

19. The promotor of the coxa (figures 4 and 5) ; a strong. two-])art muscle
extending from the anterior portion of the prepleurite and the antero-
lateral part of the tergum before the supracoxal inflection, 19a, and from
the posterior part of the prepleurite, 19b, to the tendon of the membrane
near the anterior portion of the basal coxal rim.

These basal muscles of the coxa have six cardinal group functions,
promotion, remotion, adduction, abduction, anterior rotation, and
posterior rotation. In the promotion of the coxa, muscle 19 plays
the important role; it is this muscle which extends the foreleg so
rapidly. The antagonists effecting the remotion of the coxa are 10,
11, and 13. In the adduction tlie large three-part 9 is the impor-

38—4141

594 The University Science Bulletin

tant muscle, tliough it is aided to some degree by muscle 18. The
antagonistic abduction is performed by any combined pull of 14,
15, 16, or 17. As already noted, 17 also rotates the coxa posteriorly,
though 12 is the primary posterior rotator. 12 is opposed by the
anterior rotator 11. In the movements of the coxa all of these
functions blend into one another imperceptibly by varying in-
tensities of contraction.

20. The tergopleurellar muscle of the prothorax (figures 6, 8, and 9) ; a
short, but stout, two-part muscle extending from the lateral part of the
tergum behind the supracoxal inflection to the tip of the posterior
sclerites of the pleurellar arch. Just what the significance and function
of this muscle may be is difficult to determine from this study.

21. The tergopleural muscle of the prothorax (figure 6) ; a slender muscle
extending from the lateral region of the tergum behind the supracoxal
inflection to the tip of the pleural apophysis. This muscle has little
function. It is serially homologous to the second tergopleural muscles
of the mesothorax and metathorax (51, 88, figures 29 and 30).

22. The sternopleural muscle of the prothorax (figure 6) ; a tensor muscle
composed of very short fibers holding the pleural apophysis close to the
furcal apophysis.

Within the prothoracic leg are the following muscles:

23. The extensor of the femorella (figures 7 and 15) ; a complex muscle which
pulls the well-developed extensor tendon of the membrane near the
posterior portion of the basal rim of the femorella from five points:
a, the posterior wall of the coxa; b, the pleurellar infolding; c, d, and e,
the upper lateral region of the tergum posterior to the supracoxal in-
flection. This muscle is an odd combination of numerous short bundles,
which gives the muscle a wide area of attachment upon the coxal wall,
and foin- long bands. Probably the short bundles enable the muscle to
extend the femorella and femur with rapidity, and the longer bands
furnish additional strength.

24. The median accessory extensor of the femorella (figures 11 and 20) ; a
slender muscle extending from the median distal region of the coxa to
a small tendon of the membrane near the median portion of the basal
femorellar rim, just posterior to the median coxofemorellar articulation.

25. The lateral accessory extensor of the femorella (figures 11 and 20); a
slender muscle extending from the posterior distal region of the coxa to
a small tendon of the membrane near the lateral portion of the basal
femorellar rim. These accessory extensors are practically vestigial muscles
in the prothoracic leg, since the median extensor has become such a
highly developed contractile organ. In the other legs the median and
lateral accessory extensors still play an important role.

26. The coxal flexor of the femorella (figures 11, 12, 15, and 19); a long,
strong muscle extending from the anterior wall of the coxa to the well-
developed tendon of the membrane near the anterior i)ortion of the
basal femorellar rim.

Levereault: The Carolina Mantis 595

27. The lateral accessory flexor of the femorella (figures 11, 15, and 19);
a wide, flat muscle extending from the distal lateroposterior region of the
coxa to a wide tendon of the membrane above the anterior portion of
the basal femorellar rim. In the other legs, the mesothoracic and
metathoracic, this muscle is as well developed as the median flexor. In
the twisting of the foreleg for a sagittal movement, and the elongation
of the coxa, the median flexor became developed as the chief flexor
muscle, while the lateral flexor became reduced to serve a.** an alleviator
of the strain of the chief flexor placed upon the median coxofemorellar
articulation.

28. The remotor of the femur (figure 15) ; a short, flat, two-piece muscle
extending from the ventral wall of the femorella to the lateral portion
of the basal rim of the femur. It is odd that a muscle so well developed
as this one should have so little function, for there is but little movement
posiiible between the femur and femorella.

29. The extensor of the tibia (figures 10, 13, and 18) ; a long, narrow muscle
composed of numerous short bundles extending from the dorsal wall of
the fenuu- to the long, flat tendon of the membrane near the dorsal
portion of the basal tibial rim.

30. The median accessory extensor of the tibia (figure 18) ; a vestigial
muscle extending from the dorsal wall of the femoral base to the long,
slender tendon of the membrane near the dorsal portion of the basal
tibial rim.

31. The flexor of the tibia (figures 10, 13, and 18); a powerful muscle of
complex structure extending from the ventral and side walls of the femur,
31a, and from the median wall of the femorella, 31b, to the highly-
developed flat tendon of the membrane near the ventral portion of the
basal tibial rim. This is the muscle which makes the pro thoracic tibia
and femur such an uncompromising vise mechanism.

32. The extensor of the tarsus (figures 14 and 16) ; a long, slender, fan-
shaped muscle extending from the median wall of the tibia to the
tendon of the membrane (see figure 17) near the dorsal portion of the
basal tarsal rim.

33. The flexor of the tarsus (figures 14 and 16) ; a long, slender mu.scle ex-
tending from the median and ventral walls of the tibia to the tendon of
the membrane near the ventral portion of the basal tarsal rim.

34. The flexor of the claws (figures 13, 14, 16, 17. 18, 21) ; a three-part muscle
extending from the lateroventral wall of the tibia (34a), from the dorsal
wall of the tibial base (34b), and from the laterodorsal area of the
femoral base (34c), to the long, slender tendon of the claw mechanism.
If the use of a good clearing solution to render the tarsus transparent,
and careful observation with a new Spencer research microscope may be
trusted, there are no intrinsic tarsal muscles in the Carolina mantis.

In the posterior region. of the prothorax are the muscles which
operate the hemicylinder formed by the skeleton of this thoracic
segment in a manner simulating that of a boom on a steam shovel.
Also in this region are the muscles of the first thoracic spiracle.

596 The University Science Bulletin

35. The tergal levator of the prothorax (figures 22 and 23) ; a wide, flat
muscle extending from the tendon of the membrane before the pretergite
of the mesothoracic tergum to the posterior lateral wall of the prothoracic
tergum.

36. The sternal levator of the i)rothorax (figures 22, 23, 25) ; a stout mu.scle
extending from the tendon of the membrane before the pretergital
apophysis of the mesothoracic tergum to the posterior lateral region of
the prothoracic sternum.

37. The tergal protractor of the prothorax (figures 22 and 23) ; a thick,
short muscle extending from the tip of the pretergital apophysis of the
mesothoracic tergum to the posterior median area of the prothoracic
tergum.

38. The sternal protractor of the prothorax (figures 22 and 25) ; a strong
muscle extending from the tip of the pretergital apophysis of the
mesothoracic tergum to the posterior lateral region of the prothoracic
sternum.

The levitation of the prothorax is produced by the combined ac-
tion of the sternal levators, the tergal levators, and the sternal
protractors. The sternal levators probably begin the lifting since
they are in advantageous positions when the prothorax is low; how-
ever, the sternal protractors may also start the levitation. Obviously
these are made by the pairs of these muscles functioning as units,
for unequal contraction of the pairs turns the prothorax to the side
of more intense contraction, except in the case of the sternal pro-
tractor. If the right protractor contracts more intensely than the
other muscles of this region, the prothorax is turned to the left.

The antagonistic movement to levitation, the depression of the
prothorax, is produced partially by the action of the tergal protrac-
tors, partially by the secondary ventral longitudinal muscles of the
mesothorax, and partially by the weight of the prothorax, its legs,
and the head.

39. The sternal spiracular muscle (figure 22) ; a vestigial muscle tying the
membrane below the first thoracic spiracle to the posterior lateral region
of the prothoracic sternum.

40. The occlusor of the first thoracic spiracle (figure 14, Plate LXXVIII) ; a
flat band extending from the lower anterior portion of the peritreme to
the straplike occlusor process.

41. The dilator of the first thoracic spiracle (figure 14. Plate LXXVIII); a
flaring band extending from the lower portion of the peritreme to the
anterior lip of the spiracle.

The Pterothoracic Muscles

From a study of dissections of fixed specimens of the adult Caro-
lina mantis it is not possible to separate definitely the prothoracic
from the mesothoracic muscles. There are muscles extending from

Levereault: The Carolina Mantis 597

the true intersegmental region to parts of the prothoracic skeleton,
muscles extending from the same intersegmental region to parts of
the mesothoracic skeleton, and there are muscles extending between
parts of the prothoracic and parts of the mesothoracic skeleton. In
this study I am arbitrarily designating some of these muscles as
prothoracic, others as mesothoracic, but acknowledge that only care-
ful observations of an ontogenetic study will furnish evidence of
segmental identities.

42. The primary ventral longitudinal muscles of the mesothorax (figures 24,
26) ; a pair of slender, ribbonlike muscles extending from the spinal
apophysis on the prothoracic sternum to the spinal apophysis on the
metathoracic sternum. These muscles are too weak to play any important
role.

43. The median secondary ventral longitudinal muscles of the mesothorax
(figures 24, 26) ; a pair of stout muscles extending from the long tendons
of the furcal apophyses of the mesothoracic sternum to the base of the
spinal apophysis on the prothoracic sternum.

44. The lateral secondary ventral longitudinal muscles of the mesothorax
(figures 24, 26) ; a pair of strong muscles extending from the tendons
of the furcal apophyses of the mesothoracic sternum to the posterior
lateral areas of the prothoracic sternum. These secondary ventral longi-
tudinal muscles of the mesothorax are the antagonists to the sternal pro-
tractors (38) of the prothorax. They also aid in turning the prothorax
from .side to side.

45. The sternal retractor of the firet epipleurite of the mesothorax (figures
22, 25, 26) ; a stout nuiscle extending from the side of the spinal apo-
physis on the prothoracic sternum to the lower portion of the first
epipleurite.

46. The posterior tergal retractor of the first epipleurite of the mesothorax
(figures 22, 23, 26) ; a twisted muscle extending from the pretergite of
the mesothoracic tergum (figure 23) to the lower portion of the first
epipleurite.

47. The anterior tergal retractor of the first epipleurite of the mesothorax
(figures 22, 23. 26) ; a short, thick muscle extending from the interseg-
mental ridge to the upper part of the first epipleurite. The contraction
of these retractors serves to hold the epipleurite as a basal point for the
fourth blanch of the extensor of the femorella of the mesothoracic leg.

48. The tergosternal muscle of the me.sothorax (figure 27) ; a long, slender
muscle extending from the median area of the mesothoracic tergum to
the midlateral part of the mesothoracic sternum. The function of this
muscle is the compression of the mesothorax, but this nuiscle of the
Carolina mantis is too weak for such work.

49. The flexor of the mesothoracic wing (figures 29 and 31) ; a wide, flat band
extending from the upper portion of the pleural infolding to the inner
part of the dorsal sclerite of the vannal axillary.

50. The anterior tergopleural muscle of the mesothorax (figure 29) ; a wide,
flat band extending from the pleural infolding to the midlateral margin

598 The University Science Bulletin

of the tergum. This muscle aids in the deflection of the posterior basal
area of the wing. It also aids in the arching of the tergum.

51. The posterior tergopleural muscle of the mesothorax (figures 29 and 31) ;
a long, slender muscle extending from the upper portion of the pleural
apophysis to the midlateral margin of the tergum. The action of this
muscle is similar to that of the anterior tergopleural muscle.

52. The median dorsal oblique muscle of the mesothorax (figure 26) ; a strong
muscle extending from the apophysis of the intersegmental ridge on the
metathoracic tergum to the posterior median area of the mesothoracic
tergum.

53. The lateral dorsal oblique muscle of the mesothorax (figures 27 and 29) ;
a wide band extending from the lateral surface of the apophysis of the
intersegmental ridge on the metathoracic tergum to the posterior lateral
area of the mesothoracic tergimi.

54. The intermediate dorsal oblique muscle of the mesothorax (figures 26
and 27) ; a flaring muscle extending from the tendon of the membrane
before the pretergite of the metathoracic tergum to the jiosterior medial
area of the mesothoracic tergum. These dorsal oblique muscles are pri-
marily retractors when acting as paired units. If those of the right side
only contract, the mesothorax is turned to the right.

55. The tergal protractor of the mesothorax (figure 26) ; a stout muscle ex-
tending from the tip of the apophysis of the intersegmental ridge on the
metathoracic tergum to the furcal apophysis of the mesothoracic ster-
num. Though not the most efficient muscle for protraction, it does ex-
tend the mesothoracic sternum forward after the pterothorax has been
arched.

56. The pleurosternal muscle of the mesothorax (figures 24, 26, 27) ; a short,
thick muscle extending from the lower portion of the pleural apophysis
to the furcal apophysis of the mesothoracic sternum. This muscle is
primarily a tensor between the pleurum and sternum.

57. The dilator of the second thoracic spiracle (figure 29) ; a thin, flaring
muscle extending from the furcal apophysis of the mesothoracic sternum
to the lower part of the peritreme. A fingerlike invagination of the mem-
brane above the posterior portion of the basal coxal rim is attached to
the middle of this spiracular muscle. Whatever the structural significance
of this feature may be, we shall leave it to the mantis for explanation.

58. The occlusor of the second thoracic spiracle (figure 16, Plate LXXVIII) ; a
short, fan-shaped muscle pulling the lower portions of the spiracular lips
against the lower rim of the peritreme.

These muscles are muscles that work upon the trunk portion of
the mesothoracic skeleton. The following muscles, though affecting
the trunk walls, are essentially the basal leg muscles of the meso-
thorax:

60. The sternal productor of the pleurellite (figure 24) ; a flat, flaring band
extending from the infolding separating the mesothoracic prepleurite
from the basisternite to the anterior border of the pleurellite.

61. The epipleuritellar productor of the pleurellite (figures 26, 27, 29) ; a
long, slender muscle extending from the infolding of the first epipleurite
to the anterior border of the pleurellite.

Levereault: The Carolina Mantis 599

62. The tergal abductor of the pleurellite (figure 26) ; a large twinned mus-
cle extending from the anterior median area of the tergum to the tendon
of the tip of the pleurellite.

63. The accessory promoter of the coxa (figure 31); a long, .slender band ex-
tending from the first epipleurite to the anterior portion of the basal
coxal rim.

64. The promoter of the coxa (figure 31) ; a large, three-part muscle extend-
ing from the anterior area of the prepleuritc (a), from the central part
of the first epipleurite (b), from the pleural infolding (c), to the stout
tendon of the membrane near the coxal rim.

65. The remotor of the coxa (figures 26, 27, 31); a large, two-piece muscle
extending from the central area of the tergum to the strong tendon of
the membrane near the posterior lateral rim of the coxa.

66. The anterior rotator of the coxa (figure 24) ; a short, stout muscle ex-
tending from the lower anterior position of the furcal arch of the meso-
thoracic sternum to the anterior median portion of the basal coxal rim.

67. The posterior rotator of the coxa (figure 24) ; a slender band extending
from the side of the spinal apophysis on the metathoracic sternum to the
posterior lateral portion of the basal coxal rim.

68. The accessory remotor of the coxa (figure 24) ; a short, flat band extend-
ing from the tip of the furcal apophysis to the posterior portion of the
basal coxal rim.

69. The abductor of the coxa (figure 24) ; a short, stout muscle extending
from the tip of the furcal apophysis to the lateral portion of the basal
coxal rim.

70. The depressor of the mesothoracic wing (figures 29, 31) ; a wide, flat band
extending from the lateral portion of the basal coxal rim to the third
epipleurite of the mesothorax.

71. The extensor of the femorella (figures 27, 29, 33, 35); a complex four-
piece muscle extending from the median basal region of the mesothoracic
coxa (a), from the lower surface of the furcal apophysis (b), the central
area of the tergum (c), the upper part of the first epipleurite (d), to the
highly developed tendon of the membrane near the median portion of
the femorellar basal rim.

The following muscles are wholly intrinsic muscles of the meso-
thoracic leg:

72. The anterior accessory extensor of the femorella (figures 33 and 34) ; a
strong, two-piece muscle extending from the anterior basal region of the
coxa to the tendon of the membrane near the anterior mesal portion of
the femorellar base.

73. The posterior accessory extensor of the femorella (figures 33 and 34) ; a
stout muscle extending from the posterior basal region of the coxa to the
tendon of the membrane near the posterior medial portion of the femorel-
lar base.

74. The anterior flexor of the femorella (figures 34 and 35) ; a strong, two-
piece muscle extending from the basal anterior area of the coxa to the
tendon of the membrane near the anterior lateral portion of the femorel-
lar base.

75. The posterior flexor of the femorella (figures 34 and 35) ; a strong, two-

600 The University Science Bulletin

piece muscle extending from the basal lateral area of the coxa to the
tendon of the membrane near the posterior lateral portion of the femorel-
lar base.

From the condition of the extensor and flexor muscles of the
femoreHa of the mesothoracic and metathoracic legs it is apparent
that the basic plan of the musculature of the mantis coxa consists of
two flexors nearly equal in size, and each composed of two bands,
and three flexors of unequal proportions. The musculature of the
prothoracic coxa is a modification of this basic plan. The anterior
flexor has been greatly developed, and the posterior flexor reduced;
the median extensor has been well developed, but the accessory ex-
tensors are mere vestiges.

76. The femorellar reductor of the femur (figure 36) ; a .short, stout nuiscle
extending from the ventral wall of the femorellu to the tendon of the
posterior portion of the ba.sal femoral rim. Like its homologue in the
prothoracic leg, this muscle of the mesothoracic leg has practically no
function, for there is but little movement between the femorella and
femur.

77. The extensor of the tibia (figure 36) ; a long muscle composed of numer-
ous short bundles extending from the dorsal wall of the femur (77a), and
from the anterior surface of the femorella (77b— not drawn) to the long,
flat tendon of the membrane near the dorsal portion of the tibial base.

78. The accessoiy extensor of the tibia (not figured) ; looks very much like
30b of the prothoracic leg (see figure 18) ; a vestigial muscle extending
from the dorsal basal region of the femur to the long, slender tendon of
the membrane near the dorsal portion of the tibial base.

79. The flexor of the tibia (figures 36, 38) ; a long, complex muscle of numerous
short bundles and long bands, extending from the ventral and lateral walls
of the femur to the large, flat tendon of the membrane near the ventral
portion of the tibial base.

80. The extensor of the tarsus (figures 37 and 40) ; a long, slender mascle ex-
tending from the posterior wall of the tibia to the tendon of the membrane
near the dorsal portion of the tarsal base (see figure 39).

81. The flexor of the tarsus (figures 37 and 40) ; a short, flat muscle extending
from the distal ventral region of the tibia to the tendon of the membrane
of the tarsal base.

82. The unguiflexor muscle (figures 42, 40, 37,, 38, 36) ; a three-piece muscle
extending from the anterior ventral region of the tibia (82a), the dorsal
wall of the tibial base (82b), the po.st.erior basal region of the femur (82c
—see figure 36), to the long tendon of the claw retractor mechanism.

In the metathorax the following muscles work upon the skeleton
of the Semite:

83. The sternal retractor of the metathorax (figure 24); a slender, flat band
extending from the side of the spinal apophysis on the metathoracic
sternum to the anterior portion of the infolding separating the metatho-
racic prepleurite from the basisternite. This muscle is probably serially

Levereault: The Carolina Mantis 601

homologous to 45 of the mesothorax, differing from that muscle in being
reduced, and having shifted its apex from the first epipleurite of the
metathorax to the sternal region. The reduced condition indicates this
muscle has only a minor function.

84. The lateral secondary ventral longitudinal mu.scle of the metathorax
(figures 24 and 26) ; a big muscle extending from the long tendon of the
furcal apophysis of the metathoracic sternum to the posterior surface of
the furcal apophysis of the mesothoracic sternum.

85. The tergal retractor of the first epipleurite of the metathorax (figure 26) ;
a strong muscle extending from the ajiophysis of the intersegmental
ridge of the metathoracic tergum to the lower portion of the first
epipleurite.

86. The flexor of the metathoracic wing (figures 30 and 32) ; a short, flat
muscle extending from the upper portion of the pleural infolding to the
inner surface of the dorsal sclerite of the vannal axillary.

87. The anterior tergopleural muscle of the metathorax (figures 28 and 30) ;
a wide, flat band extending from the upper portion of the pleural infold-
ing to the midlateral margin of the tergum. This muscle deflects the
posterior basal area of the metathoracic wing, and also aids in arching
the tergum.

88. The posterior tergopleural muscle of the metathorax (figures 28 and 30) ;
a slender, flat muscle extending from the upper surface of the pleural
apophysis to the midlateral margin of the tergum.

89. The median dorsal oblique muscle of the metathorax (figure 26) ; a
strong, flaring muscle extending from the posterior median area of the
metathoracic tergum to the tendon of the membrane near the anterior
border of the first abdominal tergum.

90. The lateral dorsal oblique muscle of the metathorax (figures 26 and 28) ;
a wide, flat band extending from the posterior median region of the
metathoracic tergum to the anterior lateral region of the first abdominal
tergum. The.se dorsal oblique muscles of the metathorax are retractors
of the first abdominal tergum.

91. The lateral depressor of the first abdominal tergum (figure 26) ; a flaring
muscle extending from the furcal apophysis of the metathoracic sternum
to the lateral margin of the first abdominal tergum. This muscle is
probably serially homologous to 55 of the mesothorax.

92. The pleurosternal muscle of the metathorax (figures 24, 26, 28) ; a short,
thick muscle extending from the lower surface of the pleural apophysis
to the upper surface of the furcal apophysis. This is a tensor muscle.

About the leg base of the metathorax are these muscles:

93. The sternal productor of the pleurellite (figures 24 and 32) ; a flat, flaring
band extending from the lower portions of the prepleurite and basister-
nite to the anterior border of the pleurellite.

94. The epipleuritellar productor of the pleurellite (figures 28, 30, 32) ; a long,
slender muscle extending from the lower portion of the infolding of the
first epipleurite to the anterior border of the pleurellite.

95. The tergal abductor of the pleurellite (figures 26 and 32) ; a large three-
part muscle extending from the anterior median area of the tergum to the
strong tendon of the tip of the pleurellite.

602 The University Science Bulletin

96. The piomotor of the coxa (figure 32) ; a strong, three-part muscle extend-
ing from the midanterior region of the prepleurite (a), the central area
of the first epipleurite (b), the pleural ridge (c), to the large tendon of
the membrane near the anterior lateral portion of the coxal base.

97. The remotor of the coxa (figures 26. 28. 32) ; a big muscle extending from
the median area of the tergum to the large tendon of the membrane near
the posterior lateral portion of the basal coxal rim.

98. The anterior rotator of the coxa (figure 24) ; a flat muscle extending from
the anterior part of the furcal arch to the anterior portion of the basal
coxal rim.

99. The posterior rotator of the coxa (figure 24) ; a wide, flat muscle extend-
ing from the tip of the fin-cal apophysis to the posterior portion of the
basal rim of the coxa.

100. The abductor of the coxa (figure 24) ; a twisted muscle extending from
the tip of the furcal apophysis to the lateral portion of the basal coxal rim.

101. The adductor of the coxa (figure 24) ; a. flat band extending from the
furcal apophj'sis to the posterior medial portion of the coxal base.

102. The depressor of the metathoracic wing (figures 28, 30, 32) ; a wide, flat
muscle extending from the lateral portion of the basal coxal rim to the
third epipleurite of the metathorax.

103. The chief extensor of the femorella (figures 41, 26, 28, 30) ; a big, four-
part muscle extending from the median basal area of the coxa (a), the
lower surface of the furcal apophysis (b), the anterior median area of
the tergum (c), and the upper part of the first epipleurite (d), to the
highly developed tendon of the membrane near the median portion of the
femorellar base.

The pterothoracic muscles cannot be dismissed after merely in-
dicating their primary individual functions in their names, especially
those of the Carolina mantis. They form a peculiar modification
of the basic thoracic musculature. The most startling fact in this
modification is the absence of the primary dorsal longitudinal
muscles in both the mesothorax and metathorax. This absence
makes the dissector wonder how such an insect moves its wings for
flight, and the mantis does flutter, if not fly.

Since the living insect is able to use its wings, it must be that
muscles other than the primary flight muscles of the generalized
pterygotan perform the arching of the tergum necessary for the
downstroke of the hind wings. It is probable that the following
contractions arch the metathoracic tergum of the Carolina mantis;
the tergal retractors of the first cpipleurites (85) hold those two
sclerites so they serve as relatively firm bases for the epipleuritellar
branches of the femorellar extensors (103d), and when these muscles
contract they pull back upon the first cpipleurites, which stress is
transmitted through the tergal retractors (85), which act as tensors,
to the anterior border of the metathoracic tergum. When this

Levereault: The Carolina Mantis 603

border is pulled backward, the stress is relieved by an arching of
the tergum over the wing fulcra, the uppermost points of the pleural
infoldings. In this arching the anterior tergopleural muscles aid
by pulling the posterior lateral regions of the tergum downward and
forward.

The antagonistic function, the depression of the tergum which
causes the upward stroke of the wings, may be produced by the
tergal abductors of the pleurellites (.95), the tergal branches of the
femorellar extensors (103c), and the remotors of the coxae (97). In
the more generalized thorax this depression of the tergum is per-
formed by a pair of tergosternal muscles. Such muscles, though
present in the mesothorax of the Carolina mantis, are not in the
metathorax.

In comparing the condition of the musculature of the metathorax
with that of the mesothorax, in the Carolina mantis, one is surprised
in finding the musculature of the mesothorax more elaborate than
that of the metathorax, in spite of the fact that the metathoracic legs
are used to maintain the depending position so often assumed by the
mantis. There is one more pair of epipleuritellar retractors in the
mesothorax, and the sternal epipleuritellar retractors of the meta-
thorax (83) have shifted their apices to the sides of the basister-
nite. It would seem that since the metathoracic wings are the flight
wings, a more elaborate musculature about the first epipleurites
would be necessary to compensate for the indirect manner of arching
the tergum. Though the tergosternal muscles of the mesothorax are
practically valueless, it would seem that since these muscles are
present in the mesothorax they should be well-developed contractile
organs in the metathorax. The Carolina mantis is indeed a pe-
culiar animal.

The following muscles are intrinsic leg muscles of the metathorax:

104. The anterior accessory extensor of the femorella (figure 41) ; a muscle
very similar in form and serially homologous to 72 of the mesothoracic
leg.

105. The posterior accessory exten.sor of the femorella ; .similar in form and
serially homologous to 73 of the mesothoracic leg.

106. The anterior flexor of the femorella (figure 41) ; serially homologous to 74.

107. The posterior flexor of the femorella (figure 41); serially homologous
to 75.

By comparing figures 35 and 41, it may be seen that the coxa!
musculature of the mesothorax is very similar to that of the meta-
thorax. Because of this fact, I have not drawn figures of the muscles
of the telopodite of the metathoracic limb; however, this does not

604 The University Science Bulletin

mean I have not dissected this limb. The equivalents of the meso-
thoracic leg in the metathoracic are:

108. The reductor of the femorella; serially homologous to 76.

109. The extensor of the tibia; serially homologous to 77.

110. The acce.?sory extensor of the tibia; serially homologous to 78.

111. The flexor of the tibia; serially homologous to 79.

112. The extensor of the tarsus; serially homologous to 80.

113. The flexor of the tarsus; serially homologous to 81.

114. The unguiflexor muscle; serially homologous to 82.

REFERENCES
For a good bibliography on the thorax see Snodgrass, R. E., "The
Thoracic Mechanism of the Grasshopper, and its Antecedents,"
Smiths. Misc. Coll., vol. 82, No. 2, 1929.

PLATE LXX

Fig. 1. Neck muscles from median plane.
Fig. 2. Median muscles removed.
Fig. 3. Outer muscles of neck.

Levereault: The Carolina Mantis

605

PLATE LXX

FIG. 3

606 The University Science Bulletin

PLATE LXXI

Fig. 4. Anterior prothoracic muscles from median plane.

Fig. 5. Basal muscles of right foreleg.

Fig. 6. Muscles above base of right foreleg from median plane.

Fig. 7. Median muscles removed from above dissection.

Fig. 8. Muscles posterior to tergal supracoxal inflection, from outer view.

Fig. 9. Outer mu.scles above foreleg base.

I.evereault: The Carolina Maxtis

607

PLATE LXXI

Fig. 9

608 The University Science Bulletin

PLATTE LXXII

Fig. 10. Muscles in right femur, .«ide view.

Fig. 11. Cross section through distal part of right coxa.

Fig. 12. Cross section through basal part of right coxa.

Fig. 13. Cross section through middle of right femur.

Fig. 14. Cross section through distal portion of right tibia.

Fig. 15. Muscles in right coxa and femorella, side view.

Fig. 16. Mu.';cles in right tibia from side view.

Fig. 17. Tendons and articulation of right tarsal base to tibia.

Fig. 18. Top view of muscles in right femur and femorella.

Fig. 19. Flexors of right femorella.

Fig. 20. Extensors of right femorella.

Fig. 21. Median section of right tarsus.

Levereault: The Carolina Mantis

609

PLATE LXXII

<34

Fig 17

Fig 21

39—4141

610 The University Science Bulletin

PLATE LXXIII

Fig. 22. Posterior prothoracic and anterior mesothoracic

muscles from median plane.
Fig. 23. Dorsal muscles of these regions.
Fig. 24. Ventral muscles of mesothorax and metathorax-
Fig. 25. Ventral posterior prothoracic muscles.

Levereault: The Carolina Mantis

611

PLATE LXXIII

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FIG. 24

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The University Science Bulletin

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Levereault: The Carolina Mantis

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614 The University Science Bulletin

PLATE LXXV

Fig. 33. Extensors of right middle femorella.

Fig. 34. Flexors of right middle femorella.

Fig. 35. Sideview of femorellar muscles within right middle coxa.

Fig. 36. Sideview of muscles within right middle femur and

femorella.

Fig. 37. Cross section through distal part of right middle tibia.

Fig. 38. Cross section through middle of right middle femur.

Fig. 39. Tendons and articulation of right middle tarsus to tibia.

Fig. 40. Sideview of muscles in right middle tibia.

Fig. 41. Sideview of muscles in right hind coxa.

Fig. 42. Median section of middle tarsus.

Levereault: The Carolina Mantis

615

PLATE LXXV

FiC. 41

FIG. 42

616 The University Science Bllletin

PART 3— THE ABDOMINAL MUSCULATURE

After studying for some time the cephalic and thoracic muscula-
ture of this insect, one tends to accept the notion unconsciously that
the musculature occupies most of the space within the insect body.
It is somewhat surprising to start dissecting the abdomen and find
its musculature crowded closely against the skeletal walls. In this
region the dissector is made to realize that the muscles associated
with what might be termed the manufactory of the insect have
functions different from those of the transportation and accumulat-
ing units. The muscles of the pregenital portion of the Carolina
mantis abdomen indicate by their forms and attachments that res-
piration and support of the large hull are their main concerns.

The muscles of the first abdominal segment differ from those of
the second and third segments, because the first tergum is function-
ally an anchorage of the abdomen to the thorax, and the first sternum
is reduced to a functionless condition.

1. The longitudinal retractor of the second abdominal tergum (figure 4) ; a
broad, flat sheet extending from the intersegmental region of the first
tergum to the membrane before the second tergum.

2. The median oblique retractor of the second abdominal tergum (figures
2 and 4) ; a flat flaring muscle extending from tlie posterior median region
of the first tergum to the membrane before the second tergum. In the
succeeding segments the serial homologues of this muscle are functionally
protractors.

3. The intermediate oblique retractor of the second abdominal tergum
(figures 2 and 4) ; a short muscle extending from the posterior lateral
region of the first tergum to the membrane before the paratergite of the
second tergum. In the succeeding segments the serial homologues of this
muscle are lateral retractors of the terga (see muscle 18 of figures 3
and 4).

4. The lateral oblique retractor of the second abdominal tergum (figures 2
and 4) ; a flaring muscle extending from the midlateral region of the first
tergum to the membrane before the paratergite of the second tergum.

5. The flexor of the peritreme of the first abdominal spiracle (figure 2) ; a
flat band extending from the anterior border of the paratergite of the
second tergimi to the lower anterior edge of the peritreme of the first
spiracle. The .serial homologues of this muscle in the succeeding segments
are lateral sternal retractors of the terga (see muscle 20 of figures 3 and 4).

6. The retractor of the peritreme of the first abdominal spiracle (figure 2) ;
a slender muscle extending from the anterior lateral area of the first
tergimi to the lower anterior rim of the peritreme of the first spiracle.

7. The dilator of the first abdominal spiracle (figure 17) ; a thin band ex-
tending from the anterior portion of the peritreme to the dilator process
of the lower portion of the anterior spiracular lip.

Levereault: The Carolina Mantis 617

8. The occlusor of the first abdominal spiracle (figure 17) ; a stout little
muscle extending from the occlusor process of the lower jiortion of the
posterior spiracular lip to the dilator jirocess of the anterior lip.

9. The longitudinal retractor of the second abdominal sternum (figures 1
and 4) ; a flat, flaring sheet extending from the posterior surface of the
furcal apophysis of the metathoracic sternum to the anterior border of
the second abdominal sternum.

10. The oblique retractor of the second abdominal sternum (figures 1 and
4) ; a slender band extending from the posterior portion of the meta-
thoracic sternum to the anterior lateral margin of the second abdominal
sternum.

Tlic muscles in the second abdominal segment are:

11. The longitudinal retractor of the third abdominal tergum (figure 4); a
broad, flat sheet extending from the anterior region of the second tergum
to the membrane before the third tergum.

12. The median protractor of the third abdominal tergum (figure 3) ; a flat,
flaring muscle extending from the posterior median region of the second
tergum to the membrane before the third tergum.

13. The lateral protractor of the third abdominal tergum (figure 3) ; a flat,
flaring muscle extending from the posterior lateral region of the second
tergum to the anterior edge of the paratergite of the third tergum.

14. The lateral oblique retractor of the third abdominal tergum (figures 3
and 4) ; a short, flat muscle extending from the posterior lateral region
of the second tergum to the anterior edge of the paratergite of the third
tergum.

15. The anterior tergosternal muscle (figure 4) ; a cylindrical muscle extend-
ing from the midlateral region of the second tergum to the midlateral
region of the second sternum.

16. The posterior tergosternal muscle (figure 4) ; a slender, cylindrical
muscle extending from the posterior lateral region of the second tergum
to the posterior lateral region of the second sternum. These tergo-
stemals, functioning as compressors, are the important respiratory muscles
of the pregenital segments

17. The anterior lateral retractor of the second sternum (figure 4) ; a
radiating group of bundles extending from the anterior lateral region of
the second tergum to the membrane above the anterior lateral border of
the second sternum. This group of bundles keeps the lateral membrane
tucked inside the paratergite. It also aids in the compression of the
segment for respiration.

18. The posterior lateral retractor of the second sternum (figures 3 and 4) ;
a flat muscle extending from the anterior edge of the paratergite of the
third tergum to the membrane above the posterior lateral portion of the
second sternum.

19. The tergal retractor of the third sternum (figures 3 and 4) ; a flat band
extending from the posterior lateral region of the second tergum to the
membrane before the anterior lateral corner of the third sternum.

20. The sternal retractor of the third tergum (figures 3 and 4) ; a cylindrical
muscle extending from the posterior lateral region of the second sternum
to the membrane before the paratergite of the third tergum. These re-

618 The University Science Bulletin

tractor muscles keep the four sclerotized plates of this region closely to-
gether, and fold the intervening membrane in an X-fold. When the fe-
male is gravid these muscles are stretched to an extreme degree, stretched
so much that it is a wonder she can ever flatten her abdomen again, but
she does.

21. The dilator of the second abdominal spiracle; similar in form and serially
homologous to the dilator of the first abdominal spiracle.

22. The occlusor of the second abdominal spiracle; similar in form and
serially homologous to the occlusor of the first abdominal spiracle.

23. The longitudinal retractor of the third abdominal sternum (figure 4) ; a
broad, fiat sheet extending from the anterior margin of the second
sternum to the membrane before the anterior margin of the third sternum.

24. The protractor of the third abdominal sternum (figure 4) ; a wide, flat
group of bundles extending from the posterior region of the second
sternum to the membrane before the anterior margin of the third sternum.

The musculature of the third abdominal segment is typical for the
pregenital segment of the Carolina mantis. In figure 5 are the
muscles of the third segment of the male, which are similar to those
of the female shown in figure 6.

25. The longitudinal retractor of the fourth tergum (figures 5 and 6) ; three
groups of bundles acting as a single muscle, extending from the middle
of the third tergum to the membrane before the fourth tergum.

26. The protractor of the fourth tergum (figures 5 and 6) ; a short, flat band
extending from the posterior region of the third tergum to the membrane
before the fourth tergum.

27. The oblique retractor of the fourth tergum (figures 5 and 6) ; a flat band
extending from the posterior region of the third tergum to the membrane
before the anterior lateral margin of the fourth tergum.

28. The anterior tergosternal muscle (figures 5 and 6) ; a cylindrical muscle
extending from the midlateral region of the third tergum to the mid-
lateral part of the third sternum.

29. The posterior tergosternal muscle (figures 5 and 6) ; a slender, cylindrical
muscle extending from the posterior lateral region of the third tergum to
the posterior lateral region of the third sternum.

30. The anterior lateral retractor of the third sternum (figures 5 and 6) ; a
radiating group of bimdles extending from the anterior lateral region of
the third tergum to the membrane above the anterior lateral rim of the
third sternum.

31. The posterior lateral retractor of the third sternum (figures 5 and 6) ; a
short, flat band extending from the anterior border of the paratergite of
the fourth tergum to the membrane above the posterior lateral edge of
the third sternum.

32. The tergal retractor of the fourth sternum (figures 5 and 6); a slender
muscle extending from the posterior lateral region of the third tergum to
the membrane above the anterior lateral corner of the fourth sternum.

33. The sternal retractor of the fourth tergum (figures 5 and 6) ; a muscle
extending from the posterior lateral region of the third sternum to the
membrane before the paratergite of the fourth tergum.

Levereault: The Carolina Mantis 619

34. The dilator of the third abdominal spiracle (figure 19) ; a slender, fiat
band extending from the paratergite to the dilator process of the lower
portion of the anterior spiracular lip.

35. The occlusor of the third abdominal spiracle (figure 19) ; a stout little
muscle extending from the occlusor process of the posterior spiracular lip
to the dilator of the anterior lip.

36. The longitudinal retractor of the fourth sternum (figures 5 and 6) ; three
flat bands acting as a single muscle, extending from the middle of the
third sternum to the anterior border of the fourth sternimi.

37. The protractor of the fourth sternum (figures 5 and 6) ; a band extending
from the posterior region of the third sternum to the anterior border of
the fourth sternum.

Since the musculatures of the fourth, fifth, and sixth segments of
the female, and the fourth, fifth, sixth, and seventh of the male are
identical there is no need for describing them in detail. A table of
serial homology will suffice.

SEGMENTS Ill IV V

Muscles 25 38 51

26 39 52

27 40 53

28 41 54

29 42 55

30 43 56

31 44 57

32 45 58

33 46 59

34 47 60

35 48 61

36 49 62 75 85

37 50 63 76 86

In the seventh, and eighth, the genital segments, the ninth, tenth,
and eleventh, the postgenital segments of the female mantis are the
following muscles:

77. The longitudinal retractors of the eighth tergum (figures 12 and 13) ; three
flat bands functioning as a single muscle, extending from the middle of
the seventh tergum to the membrane before the eighth tergum.

78. The median oblique retractor of the eighth tergum (figure 13) ; a flat,
flaring muscle extending^ from the posterior median area of the seventh
tergum to the membrane before the eighth tergum.

79. The intermediate oblique retractor of the eighth tergum (figures 12 and
13) ; two groups of bundles extending from the posterior lateral region of
the seventh tergum to the anterior lateral margin of the eighth tergum.

80. The lateral oblique retractor of the eighth tergum (figures 12 and 13) ;
a group of short bundles extending from the posterior lateral region of
the seventh tergum to the anterior lateral margin of the paratergite of
the eighth tergum.

VI

VII (Male)

64

77

65

78

66

79

67

81

68

82

69

83

70

118

71

119

72

84

73

120

74

121

620 The University Science Bulletin

81. The anterior tergosternal muscle of the seventh segment (figure 12); a
cylindrical muscle extending from the posterior lateral region of the
seventh tergum to the anterior lateral region of the seventh sternum.

82. The posterior tergosternal muscle of the seventh segment (figure 12) ;
a slender muscle extending from the posterior lateral region of the seventh
tergum to the anterior lateral region of the seventh sternum.

83. The anterior lateral retractor of the seventh sternum (figure 12) ; a flat
band extending from the anterior lateral region of the seventh tergum to
the membrane above the anterior lateral margin of the seventh sternum.
This muscle also aids in the retraction of the seventh sternum.

84. The posterior lateral retractor of the seventh sternum (figure 13) ; a fiat
muscle extending from the upper anterior margin of the paratergite of
the eighth tergum to the membrane above the posterior lateral margin of
the seventh sternum.

85. The dilator of the seventh abdominal spiracle ; similar in form and
serially homologous to 34 of the third segment.

86. Tlie occlusor of the seventh abdominal spiracle ; similar in form and
serially homologous to 3.5 of the third segment.

87. The lateral sternal retractor of the ovipositor base (figures 7, 11, 12) ; a-
stout muscle extending from the anterior lateral corner of the seventh
stiernum to the lower tip of the united paratergites of the eighth and
ninth terga. In addition to the retraction of the ovipositor base this
muscle serves to brace the tip of the paratergites against the more distal
muscles of the ovipositor.

88. The paratergital levator of the seventh sternum (figure 11) ; a short, thick
muscle extending from the tip of the paratergites of the eighth and ninth
terga to the midlateral region of the seventh sternum.

89 The paratergital levator of the gonopod of the ninth segment (figures 7
and 11); a short, thick muscle extending from the tip of the paratergites
of the eighth and ninth terga to the outer basal rim of the dorsal col-
linea. This muscle also extends the gonopod laterally.

90. The tergal levator of the gonopod of the ninth segment (figures 7 and
13) ; a short, flat muscle extending from anterior lateral region of the
ninth tergum to the membrane near the outer basal rim of the dorsal
collinea.

91. The paratergital abductor of the ventral collinea (figure 7); a wide,
folded muscle extending from the intersegmental ridge of the united
paratergites of the eighth and ninth terga to the lower outer portion of
the basal rim of the ventral collinea.

92. The tergal abductor of the ventral collinea (figures 7. 12, 13) ; a two-
piece muscle extending from the anterior lateral region of the eighth
tergum to the lower outer portion of the basal rim of the ventral collinea.

93. The paratergital adductor of the ventral collinea (figure 7) ; a two-piece
muscle extending from the intersegmental ridge between the paratergites
of the eighth and ninth terga to the membrane near the upper inner por-
tion of the ventral collinear base.

94. The sternal abductor of the dorsal collinea (figure 7) ; a short, thick
muscle extending from the sternum and its apophysis of the ninth seg-
ment to the outer basal rim of the dorsal collinea.

Levereaitlt: The Carolina Mantis 621

95 The accessory sternal abductor of the dorsal collinea (figure 7) ; a short
muscle extending from the ninth sternum to the lower portion of the
dorsal collinear base.

96. The abductor of the median collinea (figures 7 and 9) ; a muscle extend-
ing from the lower outer portion of the basal rim of the dorsal collinea
to the inner basal wall of the median collinea.

97. The lateral muscle of the dorsal collinea (figure 8) ; a wide, flat muscle
extending from the lateral wall of the dorsal collinea to the membrane
of its inner basal rim.

98. Tile collineafer muscle of the ventral collinea (figure 10) ; a short, thick
muscle extending from the inner basal wall of the collineafer region to the
outer basal wall of the ventral collinea.

99. The sternal retractor of the genital chamber (figure 7) ; a slender, flat
band extending from the anterior border of the seventh sternum to the
anterior edge of the genital chamber.

100. The sternal protractor of the genital chamber (figure 12) ; a flat, flaring
muscle extending from the central part of the seventh sternum to the
lower wall of the genii al chamber. This part of the genital chamber is
sclerotized and has been identified as the eighth sternum. It might be
claimed that this protractor muscle is fundamentally a retractor on the
eighth sternum, but it should be noted that the muscle apex is on the
morphologically posterior border of the sclerotic plate.

101. The longitudinal protractor of the ninth tergum (figures 12 and 13);
three flat bands functioning as a single muscle, extending from the an-
terior border of the eighth tergum to the anterior border of the ninth
tergum.

102. The median oblique retractor of the ninth tergum (figure 13) ; a wide,
flat band extending from the anterior median area of the eighth tergum
to the anterior lateral margin of the ninth tergum.

103. The intermediate oblique retractor of the ninth tergum (figures 12 and
13); a flat muscle extending from the anterior lateral region of the eighth
tergiuii to the anterior lateral margin of the ninth tergum.

104. The lateral oblique retractor of the ninth tergum (figures 12 and 13) ; a
flat muscle extending from the midlateral region of the eighth tergum to
the anterior lateral margin of the ninth tergum.

105. The dilator of the eighth abdominal spiracle; similar in form and
serially homologous to 34 of the third segment.

lOo. The occlusor of the eighth abdominal spiracle; similar in form and s<Mially
homologous to 35 of the third segment.

107. The longitudinal protractor of the tenth tergum (figures 12 and 13) ;
three flat bands functioning as a single muscle, extending from the an-
terior border of the ninth tergum to the anterior border of the tenth
tergimi.

108 The median oblique retractor of the tenth tergum (figures 13 and 15) ; a
flat, flaring muscle extending from the anterior median border of the
ninth tergum to the anterior lateral mai'gin of the tenth tergum.

109. The lateral oblique retractor of the tenth tergum (figure 15); a twisted
band extending from the anterior lateral border of the ninth tergvmi to
the anterior lateral corner of the tenth tergum.

622 The University Science Bulletin

110. The lateral retractor of the paraproct (figures 12 and 13) ; a long, flat
muscle extending from the anterior lateral border of the ninth tergum
to the lower basal rim of the paraproct.

111. The longitudinal retractor of the eleventh tergum (figures 12, 13, 15); a
slender band extending from the anterior median border of the tenth
tergum to the anterior border of the eleventh tergum.

112. The median retractor of the paraproct (figures 13 and 15) ; a two-piece
muscle extending from the anterior median region of the tenth tergum to
the upper median border of the paraproct.

113. The dorsal flexor of the cercus (figure 15) ; a flat, flaring muscle extending
from the anterior median area of the tenth tergum to the inner dorsal
portion of the cereal base.

114. The ventral flexor of the cercus (figure 15) ; a flaring band extending
from the anterior border of the tenth tergum to the inner ventral portion
of the cereal base.

115. The dorsal extensor of the cercus (figure 15) ; a flaring band extending
from the anterior lateral region of the tenth tergum to the outer dorsal
portion of the cereal base.

116. The ventral extensor of the cercus (figure 15) ; a short muscle extending
from the curled lateral portion of the tenth tergum to the outer ventral
portion of the cereal base.

117. The median retractor of the paraproct (figures 12 and 13); a flat band
extending from the lower base of the paraproct to the lower distal region
of the paraproct.

The rectal dilator muscles are not included in this study because
the specimens available were not fixed well enough for the dissection
of such muscle bundles.

In the eighth, ninth, tenth, and eleventh segments of the male are
some muscles which are obviously serially homologous to those in
the posterior segments of the female, some which are probably
strongly modified homologues, and others which are special male
developments.

101. The longitudinal retractor of the ninth tergum (figure 18); three flat
bands functioning as a single muscle extending from the anterior region
of the eighth tergum to the membrane before the anterior border of the
ninth tergum. Homologous to 101 of the female.

102. The oblique protractor of the ninth tergum (figure 18) ; a flat, flaring
band extending from the posterior median region of the eighth tergum to
the membrane before the anterior border of the ninth tergum. Homol-
ogous to 102 of the female.

103. The intermediate oblique retractor of the ninth tergum (figure 18); a flat
band extending from the posterior lateral region of the eighth tergum
to the anterior lateral margin of the ninth tergum. Homologous to 103
of the female.

104. The lateral oblique retractor of the ninth tergum (figure 18) ; a flat band
extending from the posterior lateral region of the eighth tergum to the
anterior lateral margin of the ninth tergum. Homologous to 104 of the
female.

Levereault: The Carolina Mantis 623

122. The anterior tergosternal muscle of the eighth segment (figure 18) ; a
stout muscle extending from the midlateral region of the eighth tergum to
the midlateral region of the eighth sternum. Probably homologous to 92
of the female.

123. The sternal retractor of the ninth tergum (figure IS) ; an oblique muscle
extending from the posterior lateral region of the eighth sternum to the
anterior lateral corner of the ninth tergum.

124. The longitudinal retractor of the ninth sternum (figure 18) ; a flat band
extending from the anterior border of the eighth sternum to the anterior
border of the ninth sternum.

125. The median protractor of the ninth sternum (figure 18) ; a flat band ex-
tending from the posterior median region of the eighth sternum to the
anterior median border of the ninth sternum.

126. The lateral protractor of the ninth sternum (figure 18) ; a cylindrical
muscle extending from the posterior median region of the eighth sternum
to the anterior lateral corner of the ninth sternum. This muscle also
twists the sternum.

107. The longitudinal retractor of the tenth tergum (figure 18) ; three bands
functioning as a single muscle extending from the anterior border of the
ninth tergum to the anterior border of the tenth tergum. Homologous to
107 of the female.

108. The oblique protractor of the tenth tergum (figure 18) ; a long, flat band
extending from the posterior median region of the ninth tergum to the
anterior lateral border of the tenth tergum. Homologous to 108 of the
female.

109. The lateral oblique retractor of the tenth tergum (figure 18) ; a muscle
extending from the posterior lateral region of the ninth tergum to the
curled anterior lateral portion of the tenth tergum. Homologous to 109
of the female.

127. The tergal retractor of the ninth sternum (figure 18) ; a thick, flat band
extending from the lateral region of the ninth tergum to the anterior
lateral edge of the ninth sternum.

128. The sternal retractor of the intersternal membrane (figure 18) ; a wide,
flat band extending from the anterior lateral edge of the ninth sternum to
the intersternal membrane above the ninth sternum.

129. The left paraproctal muscle of the right lobe of the male genitalia (figure
23) ; a flat band extending from the lower border of the left paraproct to
the median process of the right genital lobe.

130. The right paraproctal muscle of the right genital lobe (figure 23) ;
a flat band extending from the lower border of the right paraproct to the
median process of the right genital lobe.

131. The median retractor of the v^entral membrane of the right genital lobe*
(figure 23) ; a wide, flat band extending from the median process to the
middle of the ventral membrane of the right genital lobe.

132. The distal flexor of the right genital lobe (figure 23) ; a cylindrical muscle
extending from the middle of the lateral process of the right genital lobe
to the upper surface of its distal cap.

133. The basal retractor of the right genital lobe (figure 23) ; a cylindrical

* The muscles of the genital lobes of the male are not paired muscles.

624 The University Science Bulletin

imif^clp extending from the middle of the hiteral process of the right geni-
tal lobe to the anterior median border of the ninth sternum.

134. The retractor of the right genital lobe (figure 23) ; a cylindrical muscle

extending from the base of the ventral sclerite of the median lobe to the
ba*?al lip of the lateral process of the right lobe.

135. The basal flexor of the right genital lobe (figures 23 antl 24) ; a flat
muscle extending from the inner surface of the lateral process of the
right genital lobe to the midventral region of the lobe.

136. The dorsoventral nuiscle of the right genital lobe (figure 24) ; a ;-mall
flaring muscle extending from the upper surface of the midlateral region
of the lobe to the crescenteric sclerite in the midventral region of the
lobe.

137. The interlobular mu.scle between the right and left genital lobes (figure
22) ; a flat, flaring band extending from the tip of the median process of
the right lobe to the median apophysis of the left lobe.

138. The apophyseal tensor of the left genital lobe (figure 22) ; a wedge-shaped

nuiscle extending from the median apophysis of the left lobe to the
median portion of the lateral sclerite of the left lobe.

139. The flexor of the lateial sclerite of the left genital lobe (figures 21 and
22) ; a slender muscle extending from the median portion of the lateral
sclerite to the ventral surface of the earlike lobe of the median genital
lobe.

140. The flexor of the median apophj'sis of the left lobe (figiu'e 21) ; a large
muscle extending from the lateral basal region of the ventral sclerite of
the left lobe to the base of its median apophysis.

141. The retractor of the ventral sclerite of the left lobe (figures 20 and 21) ;
a stout muscle extending from the basal region of the ventral sclerite of
the median lobe to the lower surface of the lateral basal region of the
ventral sclerite of the left lobe.

142. The median depressor of the left lobe (figures 20 and 21) ; a curved, flat
band extending from the midventral region of the ventral sclerite of the
median genital lobe to the inner basal sclerite of the left lobe.

143. The retractor of the tip of the median genital lobe (figure 20) ; a large,
stout muscle extending from the middle of the ventral sclerite of the
median lobe to the membrane before the tip.

144. The protractor of the median genital lobe (figure 21) ; a large muscle ex-

tending from the anterior median region of the ninth sternum to the
basal rim of the median lobe.

I shall not defend the names of these genital muscles very strongly,
for it would take more observation of the movements of the genital
lobes than I have made to interpret the functions of their muscles.
In the tenth segment of the male are tlie muscles of the eleventh
tergum, the paraprocts, and the cerci, all similar to those of the
female:

111. The longitudinal retractor of the eleventh tergum (figure 18); a slender,
flat band extending from the anterior border of the tenth tergum to the
anterior border of the eleventh. Homologous to 111 of the female.

Levereault: The Carolina Mantis 625

112. The median retractor of the paraproct (figure 18) ; a single band extend-
ing from the anterior border of the tenth tergum to the upper median
border of the paraproct. Homologous to 112 of the female.

113. The dorsal flexor of the cereus (figure 18) ; simihir in form and homol-
ogous to 113 of the female.

114. The ventral flexor of the cercus (figure 18); similar in form and homol-
ogous to 114 of the female.

115. The dorsal extensor of the cercus; similar in form and homologous to 115
of the female.

116. The ventral extensor of the cercus; similar in form and homologous to

116 of the female.
110. The lateral retractor of the paraproct (figure 18) ; similar in form and
homologous to 110 of the female.

117. The median retractor of the paraproct (figure 18) ; similar in form and
homologous to 117 of the female.

REFERENCES

See Snodgrass, R. E., ''The Abdominal Mechanisms of a Grass-
hopper/' Smiths. Misc. Coll., vol. 94, No. 6, 1935.

40—4141

626 The University Science Bulletin

PLATE LXXVI

Fig. 1. Ventral muscles of second abdominal sternum.

Fig. 2. Lateral view of muscles of first abdominal tergum.

Fig. 3. Dorsal and lateral muscles between second and third segments.

Fig. 4. Lateral view of muscles of first and second segments.

Fig. 5. Lateral view of musculature of male third segment.

Fio. 6. Lateral view of musculature of female third segment.

Levereailt: The Carolina Mantis

627

PLATE LXXVI

Fig I

Fig. 4

Fig. 3

Fig 6

628 The University Science Bulletin

PLATE LXXVII

Angular dorsal view of basal ovipositor muscles.
Side view of basal muscles of right dorsal collinea.
Side view of abductor of right median collinea.
Side view of collineafer muscle of right ventral collinea.
Angular dorsal view of ovipositor base with upper muscles
removed.
Fig. 12. Lateral view of terminal segmental muscles of the female.

Fig.

7,

Fig.

8.

Fig.

9.

Fig.

10.

Fig.

11.

Levereault: The Carolina Mantis

629

PLATE LXXVII

Fig. 10

Fig. 7

VII.

vw.

^951

FIG. II

FIG. 12

630 The University Science Bulletin

PLATE LXXVIII

Fig. 13. Muscles of seventh, eighth, ninth, tenth, and eleventh terga of
female.

Fig. 14. Muscles of first thoracic spiracle.

Fig. 15. Enlarged view of muscles of ninth, tenth, and eleventh terga of
female.

Fig. 16. Muscles of second thoracic spiracle.

Fig. 17. Muscles of first abdominal spiracle.

Fig. 18. Lateral view of terminal segmental muscles of male.

Fig. 19. Muscles of third abdominal spiracle.

Levereal'lt: The Carolina Mantis

631

PLATE LXXVni

Fig. 15

Fig. 16

VIII

FIG. 18

Fig. 19

632 The University Science Bulletin

PLATE LXXIX

Fig. 20. Muscles on median lobe of male genitalia.

Fig. 21. Muscles within left lobe of male genitalia.

Fig. 22. Muscles at base of left lobe of male genitalia.

Fig. 23. Muscles of right lobe of male genitalia.

Fio. 24. Lateral muscles in right lobe of male genitalia.

Levereault: The Carolina Mantis

633

PLATE LXXIX

FIG. 20

129

FIG. 23

Fig 21

FIG. 22

Fig. 24

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXV] June 1, 1938 [No. 22

A Monographic Revision of the North American Species
of Stenehnis (Dryopidae: Coleoptera)*

MILTON W. SANDERSON,

Deiiartinent of Entomology. Univei>i1y of Kansas

Absthact: Thi.s paper, according to iti< title, is a moncgrajihic revi.'^ion of
the North American species of Stenelmi.s. It contain.s virtually all that has
been written on thi.'^ genu^; of Dryopidae for the Western Hemisphere. This
includes all that is known of the biology and morphology of the genus in
addition to the taxonomy.

Approximately 600 species of Dryopidae have been described, of which
sixty-eight belong to the genus Stenelmis. Fifteen species of this number
have been described from North and South America, of which four species:
Stenelmis sidcata Blatch. {blatchlcyi Musgr.), S. linearis Zimm., S. sordida
Mots., and S. elongata Mots, are reduced to synonymy in this paper. Eight-
een species of Stenelmis are described here for the fir.^t time. They are:
Stenelmis sexlineata, S. exigua, S. beameri, S. lateralis, S. concinna, S. tarsalis,
S. knoheli, S. exilis, S. niera, S. douglasensis, S. grossa, S. parva, S. hungerfordi,
S. mii-abiUs, S. antennnlis, S. musgravei, S. decorata, and S. convexida.

With the exception of Stenelmis crenata (Say), the types, cotypes, or
l>aratypes have been examined for each of the American species. Since the
type or types of S. crenata are lost, new types are designated.

The key to Stenelmis is based to some extent upon the color patterns of
the elytra. Considerable use has, however, been made of the tarsal claws
and segments, tubercles of the pronotum, apical abdominal emargination,
granulation, the male genitalia, and a number of other characters not here-
tofore used. Males have been studied of all the species except two. With
these exceptions, a figure of the genitalia is given for each species. This is
the first time that a comparative study has been attempted of the male
genitalia of Stenelmis. Technicjue is given for dissecting and mounting the
genitalia.

A redescription or notes have been given for each of the American species.

A brief history of the classification of the entire family has been given
from the time that the first Dryopid was described. The family was un-
known to Linnaeus.

Although no fo.ssil Stenelmis are known, a geological history of the family
is given.

* Submitted to the Department of Entomology and the Facvdty of the Graduate School
of the University of Kansas in partial fulfillment of the requirements for the degree of doctor
of philosophy.

(635)

636 The 'University Science Bulletin

TABLE OF CONTENTS

PA8B

Introduction 637

Acknowledgments 637

Historical Review of Stenelmis in North America 638

History of Classification of Family 639

Family Characteristics 642

Biology and Morphology 643

Need for Research 643

Life History 643

Respiration and Habits of the Larva 644

Respiration and Habits of the Adult 646

Description of the Larva 649

Description of the Pupa 652

Geographical Distribution 652

Geological Distribution 653

Bibliography 654

Bibliography of Genus 654

Genotype 654

Description of the Genus Stenelmis 654

Secondary Sexual Characters of Stenelmis 655

Specific Characters in Stenelmis 655

The Male Genitalia of Stenelmis 658

Technique for Dissecting Male Genitalia 659

Systematic Treatment 660

Key to Genera of Dryopidae of the United States 660

Key to Groups of Stenelmis 661

Nubifera Group 661

Key to the Species of the Crenata Group 663

Key to the Species of the Sinuata-Humerosa Group 684

Addenda 706

Stenelmis geayi Grouv 706

Stenelmis nevermanni Hntn 709

Cylloepus piincticollis (Hntn.) 711

Micro cylloepus pusillus (Lee.) 711

Stenelmis canaliculata (Gyll.) 711

List of American Species of Stenelmis 711

Literature Cited 712

Plates 715, 717

Sanderson: Species of Stenelmis 637

INTRODUCTION

SEVERAL years ago when I attempted to determine the material
in the Dryopid genus Stenelmis in the Francis Huntington Snow
collection at the University of Kansas, it became apparent that
practically nothing could be done, M'ith the present literature, to-
ward the identification of our forms. Nearly all of the descriptions
are brief, important structural characters were overlooked by early
workers, and as a result the specimens of Stenelmis in most col-
lections have been misidentified or set aside without names. Added
to this difficulty, the specimens are usually clothed with an in-
crustation of dirt or other debris which has further rendered identi-
fication almost impossible. Thus, it became necessary to examine
the types of our species in order to approach any accuracy of
determination. With the exception of one, Stenelmis crenata (Say),
the types have been studied.

No monographic studies have been attempted previously although
LeConte (1852) and Horn (1870) brought our species together in
synoptic arrangement. The purpose of this paper is to redescribe
all known North American species of Stenelmis, to bring the scanty
and scattered literature together, and to make known a number of
new and interesting forms. Keys and illustrations are presented
which, it is hoped, will aid in the identification of our species. For
the sake of completeness, descriptions of Stenelmis geayi Grouv.
and Stenelmis nevermanni Hntn., the only species known in the
Western Hemisphere outside of the United States, are appended at
the end of this paper.

Acknow^ledgments

This opportunity is taken to express my deepest appreciation to
Dr. H. B. Hungerford for the encouragement and support which
have made the writing and publication of this paper possible. His
readiness in aiding me to secure materials, to examine various types
in the older collections, and to advise me on many problems under
consideration, has been a constant stimulus. I am also greatly in-
debted to Dr. Paul N. Musgrave, Fairmont, W". Va., who submitted
his large collection of Stenelmis for study, and kindly lent me much
of the literature from his extensive library on Dryopoidea. His
opinions on various questions relative to Dryopidae have been very
valuable, and I take this opportunity to express my appreciation.
To Dr. R. H. Beamer, of the University of Kansas, I owe my thanks
for the large numbers of specimens of many interesting species
which he has secured on his surveys over the United States.

638 The University Science Bulletin

Through the kindness of Dr. B. S. Cusin, of the Zoological Museum
of Moscow, U. S. S. R., I have been permitted to examine the types
of the species of Stenelmis described by Motschulsky (1859) from
North America.*

To Mr. Guy Colas, of the National Museum at Paris, we are
indebted for two specimens from the type series of Stenelmis geayi
Grouv. And to Mr. Howard E. Hinton, of Berkeley, Gal, I am
grateful for a paratype of Stenelmis nevermanni Hntn.

To the following I am indebted for the loan of material: Uni-
versity of Minnesota ; Canadian National Museum ; Warren Knaus,
McPherson, Kan.; Kansas State College; C. A. Frost, Framingham,
Mass.; 0. L. Cartwright, Clemson College, S. C; P. W. Fattig,
Emory University, Ga.; Cornell University, Ithaca, N. Y.; Dr.
H. H. Knight, Iowa State College.

I am grateful for the privilege of studying the collections of Prof.
W. S. Blatchley, Indianapolis, Ind. ; the collections of the Museum
of Comparative Zoology, Harvard College; and the Horn collec-
tion at the Philadelphia Academy of Sciences.

To Mr. H. C. Fall, Tyngsboro, Mass.; Dr. Lyman S. Henderson,
my fellow student; to Mr. C. W. Sabrosky, East Lansing, Mich.;
and to all others who have aided in this project, I wish to give my
sincere thanks.

The location of types is indicated after the description of each
species.

Historical Review of Stenelmis in North America

The genus Stenelmis was first proposed by Leon Dufour in 1835
(Add. des Sci. Nat. Second Serie, tome III, p. 158) for the reception
of Elmis canaliculata Gyll., a species described in 1808 from Europe.
Later Elmis crenata Say, described in 1924 from Pennsylvania,
was transferred to this genus. In 1852 LeConte named two more
species in the United States, S. sinuata, and S. hicarinata. In 1859
Motschulsky described S. humerosa, S. mdrkelii, S. sordida, and S.
elongata. In 1869 Zimmerman added S. linearis and S. vittipennis.
S. quadrimacidata Horn was described in 1870, at which time its
author indicated the possible synonymy of »S. linearis Zimm. with
S. humerosa Mots, in the following statement: "This may be the
species described by Motschulsky under the name of humerosa,
and, if so, the latter should have priority. The descriptions of this
and others are so vague as to leave considerable doubt of their

* Stenelmix sordida, S. elongata, S. mdrkelii, and S. humerosa.

Sanderson: Species of Stenelmis 639

identity." Tliis conclusion, while correct, had not been verified
until the present paper. In 1901 .S. nubijera Fall was described and
recorded as the first species of the genus from the Pacific coast.
In 1910 Blatchley described S. sulcata, and in 1925, S. fuscata. In
1933 Musgrave discovered that the name sulcata had been used by
Grouvelle (Not. Leyd. Mus. XIV, 1892, p. 188) for a species from
Sumatra, and proposed the name blatchleyi for this species.

History of Classification of Family

The following account is intended to give a brief history of the
important changes in the family since the first species was described.
Most of the information, until 1852, was taken from Mulsant and
Key (Hist. nat. col. Fr. 1872), and Zaitzev's account of the Dry-
opidae (Coleopterorum Catalogue, Vol. 14). The family was un-
known to Linnaeus.

1785. Geoffrey (Foure. Ent. Paris, I, p. 20) was the first to describe a species
of Dryopidae, Dermestes auriculatus, which he placed with the Der-
m est ids.

1787. Fabricius (Mantissa Insectoriim) described a species which he called
Elater dermestoides.

1791. Olivier (Encycl. method. VI, p. 298) proi)Osed the generic name of
Dryops for the species which Geoffroy had placed with the Dermestidae.

1792. Fabricius (Ent. Sj'st. I, p. 245) removed the species which he had
described as an Elaterid and made it type of the genus Parnus. At the
same time he described Pitnius prolifericornis.

1793. Panzer (Fn. Germ. 7, p. 4) described a species which he called Dyliscus
volckmnri.

1798. Latreille (Bull. Soc. Philom. I-II) described the genus Elmis.

1802. Marsham (Entom. Brit. I, i). 192) placed the same insect in the genus
Chrysomela.

1804. deBrives (Hist. Nat. IX, p. 227) associated Elmis and Dryops with the
Byrrhidae, and also placed the Heteroceridae with the same family.
Altogether they formed the group Ripicoles. He stated that Dryops
and Gyrinids made up the family Otiophores.

1806. Midler (Illig. Mag. V, p. 184) adopted the name Limnius.

1807. deBrives (Gen. Crust. II, p. 48) preserved the family Otiophores.

1811. Germar (N. Schriften Nat. Ges. Halle 1, VI, p. 41) established the genus
Potamophilus for Pamuf: acumiiiatus Fab.

1817. Leach (Cuv. Regne Anim. Ill, ]>. 268) changed the name Potamophilus
to Hydera. This genus, Dryops, and Heterocerus form the second sec-
tion of the family of Clavicornes.

1817. Leach (Zool. Misc. Ill, p. 88) placed Dryops with the Parnidea, and in

addition Parnus and Potamophilus are placed in the same group.
1825. MacLeay (Annul. Javan. ed. I, p. 34) recognized the family Parnidae.

640 The University Science Bulletin

1828. Stephens (111. Brit, Ent. II, p. 102) divided the second section of the
Clavicornes of Latreille into three families: Heteroceridae, Parnidae
and Limnidae.

1839. Westwood (Intro, to the Modern Classification of Insects, Vols. I and
II) adopted the divisions of Macleay. He called the Limnidae of
Stephens a subfamily of Parnidae, but preferred the subfamily name
of Elmidae.

1845. Redtenbacher (Die Gattimgen der deutschen Kaferf., Wien.) placed
the Parnidae, Elmidae, and Heteroceridae after the Hydrophilidae. He
placed the Georyssidae between the Dermestids and Byrrhids.

1847. Erichson (Naturg. Ins. Deutschl. Ill, p. 509) reunited all described
Dryopids into a single family, the Parnidae. This was divided into two
tribes, Unciferes and Diversicornes.

1852. LeConte divided the United States representatives of the family into
three groups: Eurypalpini, which included the genus Eurypalpus
(Psephenus) ; Dryopini, which included lara, Lutrochus, Pelonomus,
and Helichus ; and the Elmini, which included Limnius, Elmis, Stenelmis,
Macronychus and Ancyronyx.

1861. LeConte, in his Classification of the Coleoptera of North America
(Smith. Miscell. Coll. 1861), considered the family Parnidae as con-
taining three distinct subfamilies: Psephenidae, Parnidae (genuini)
which contained the tribes Larini and Pamini, and the Elmidae.

1872. Mulsant and Rey (Hist. nat. col. Fr. XXII) retained the tribes
Unciferes and Diversicornes of Erichson.

1870. Horn (Trans. Amer. Ent. Soc. Ill) recognized the three subfamilies of

LeConte.
1883. LeConte and Horn (Smith. Miscell. Coll. 507), in their Classification

of Coleoptera, recognized the subfamilies Psepheninae, Parninae, and

Elminae.

1900. Lameere (Ann. Soc. Ent. Belg. XLIV, 363) proposed to unite Psephenus
and also the Heteroceridae and Georyssidae with the Dryopidae. The
family then was divided into the following subfamilies: Psepheninae,
Parninae (including Heteroceridae), and Elmidinae (including Georys-
sidae).

1904. Ganglbauer (Die Kafer von Mitteleuropa) has used the subfamilies
Dryopinae and Helminthinae.

1908. Zaitzev recognized the subfamilies Psephenini, Dryopini, and Helminthini.

1910. Zaitzev places the species in the Psepheninae, Dryopinae, and Helminae.

1910. Blatchley (Coleoptera of Indiana, p. 677) follows LeConte and Horn,
recognizing the subfamilies Psepheninae, Parninae, and Elminae of the
family Parnidae.

1920. Leng (Catalogue of Coleoptera) considered that Psephenus should con-
stitute the family Psephenidae. The subfamily Parninae, or Parnidae
of some authors, is raised to the family rank of Dryopidae, and the
Elminae is raised to family rank, but is designated Helmidae.

1926. Forbes (Jr. N. Y. Ent. Soc. XXXIV, pp. 107-108) on a basis of wing
folding patterns of Coleoptera recognizes the superfamily Dryopoidea

Sanderson: Species of Stenelmis 641

as constituting the Ptilodactylidae, Drj'opidae (with Psephenus and
Elmis), Chelonariidae, Heteroceridae, Byrrhidae (except Nosodendron)
and the Mycetophagidae.

1927. Barthe (Miscellanea Entomologica, XXX, p. 4) follows Ganglbauer in

the naming of subfamilies.
1927. West (Ann. Ent. Soc. Amer., XXII, p. 691), although acknowledging

the usage of Psephenidae, Dryopidae, and Elmidae by Leng, preserves

Psepheninae, Dryopinae, and Elminae as subfamilies of Dryopidae.

His assumptions were based upon a study of the larvae.
1929. Carter and Zeck (Australian Zoologist, VI, p. 51), in their Monograph

of Australian Dryopidae, recognize the Psepheninae, Dryopinae, and

Helminae.

1929. Boving (Bull. Brook. Ent. Soc, 24, p. 55) studied the larva of Lara and
as a result concluded that it should constitute a separate family from
the Dryopidae. This he named the Larridae. The genus Eubrianax,
formerly included in the Dascyllidae, is placed with the Psephenidae.
Other families, in addition to the Dryopidae, were Chelonariidae,
Ptilodactylidae, and Psephenidae.

1930. Boving and Craighead (Entomologica Americana, XI, pp. 1-351) gave
us our more recent general classification of the larvae of Coleoptera.
Based upon a study of larval characters they recognized the series
Dryopoidea under which were included the following families : Ptilo-
dactylidae, Eiuypogonidae, Psephenidae, Chelonariidae and Dryopidae.
In this proposed classification the Heteroceridae is removed to the
series Dascilloidea. The genus Eurypogon is removed from the Ptilo-
dactylidae and given the rank of family. The Psephenidae is divided
into the Psepheninae and the Eubrianacinae, the latter formerly having
been included in the Dascyllidae. The Dryopidae included the sub-
families Larinae, Pelonominae (containing Pelonomus, Helichus and
Psephenoides), and Helminae (containing Dryops, Helmis, Limnius,
Ancyronyx, etc.) . The Ptilodactylidae, Eurypogonidae, and Chelonariidae
were formerlj^ included in the Dascilloidea.

1930. Blackwelder (Pan. Pac. Ent., VI) also favors the placing of Eubrianax
with the Psephenidae.

1930. Bradley (Manual of the Genera of Beetles of America North of Mexico)
included the Psephenidae, Laridae, Drj'opidae, Georyssidae, Chelonarii-
dae, and Ptilodactylidae in the superfamily Dryopoidea. He mentioned
that the family name Laridae is preoccupied by the family of sea gulls,
though based on a differently spelled generic name.

1935. Hinton (Stylops, Vol. 4, p. 173) recognizes three subfamilies, Larinae,
Dryopinae, and Elminae. He also considers the Psephenidae as a
separate family.

1936. Darlington (Psyche,- XLIII, pp. 65-83), in a paper on West Indian
Dryopidae, recognized the subfamilies Psepheninae, Dryopinae, and
Helminae.

1936. Hinton (Annals and Magazine of Natural History, Ser. 10, Vol. XVIII,
p. 89) considers the same subfamilies as in his paper of 1935.

41—4141

642 The University Science Bulletin

The generic names Elmis and Helmis have been used interchange-
ably since Hehiiis first was designated. Bedel (Ann. Soc. Ent. Fr.,
V, 1878, p. LXXV) added the "h" to Elmis, claiming that Latreille,
being French, did not use the aspirate. Since the original trans-
literation was Elmis, there is no reason why the name Helmis should
be used.

Musgrave (Proc. Ent. Soc. Wash., 37, 1935, pp. 137-138), in a
discussion of Helichus, gives a brief review of the usage of Parnus
and Dryops, and indicates why the name Parnus never should have
been used. According to this author, "Helichus being very close
to Dryops Olivier (Parnus Fabricius) was confused with the latter
genus for more than a hundred years; and Erichson in his key to
the Dryopini, which accompanied his original description, made the
mistake of separating the two genera on approximate or distant hind
coxae. . . . This error was noted by Sharp (Biol. Centr. Amer.
I, Pt. 2, 1882, p. 120) , who placed species in the genus Dryops Leach,
making Helichus a synonym. Much of the confusion was caused
by the fact that both Olivier (Encycl. method. VI, 1791, p. 297)
and Leach (Zool. Misc. HI, 1817, p. 88) described genera, applying
the name Dryops. Dryops Oliv. (Parnus Fab.) is the correct name
and is the true Dryops. The Dryops of Leach is the Helichus of
Erichson and cannot be used because the name was previously used
by Olivier. This fact validates Helichus Erichson, although the
name did not appear until 1847. Parnus Fabricius (Ent. syst. I, 1,
1792, p. 245) is a synonym of Dryops Oliv.; and, therefore, species
placed in Parnus belong to Dryops Oliv."

Family Characteristics

Dryopidae are usually characterized by being small, aquatic or
subaciuatic beetles, rarely over eight to ten millimeters in length,
and having the following characters: head usually in part retractile
and nearly always protected beneath by the prosternal lobe. In
most species this lobe, while the head is in repose, meets the labrum
and completely conceals the mouthparts. Antennae 7-11 segmented,
the great majority of the species having the latter number. Pro-
sternum generally with a posterior median process which fits into
or against a groove on the mesosternum. Coxae usually widely
separated, the anterior ones widely open behind. Hind coxae trans-
verse and usually partly protecting the hind femora. Abdomen
distinctly 5-segmented, the first few segments firmly united. Legs
usually long and slender; tarsi 5-segmented and with the last
segment often as long as the four preceding combined. Claws \'ery
long and generally robust.

Sanderson: Species of Stenelmis 643

BIOLOGY AND MORPHOLOGY
Need for Research

An examination of the literature on the biology and morphology
of Dryopidae indicates that much remains to be discovered. To
my knowledge the complete life cycle of none of the species has
been worked out, although we have considerable information on the
development of the closely related family, Psephenidae. The larvae
of but few of our species are known, the pupae of but two or three,
and the eggs of one or two. Nothing has been done on the embry-
ology. Only recently has anything been discovered regarding the
tracheation of the larva, and information on the digestive and other
systems is almost entirely lacking.

In 1835, at the time the genus Stenelmis was described, some in-
formation was given on the morphology of the adult, but nothing
was known of the biology. Brocher (1912) has added some data
on adult respiration, and suggests that this subject needs to be more
fully developed.

The larvae and adults can be found in abundance in their natural
habitat, so cjuantitics of material may be had for study. The eggs
of Stenelmis have been seen in the field once or twice, but I have
obtained them only by the dissection of adult females.

Life History

Matheson (1914) was the first to record data on the immature
stages of any of our species of Stenelmis. He found the full-grown
larvae and pupae of Stenelmis bicarinata Lee. (latitude of Truro,
Nova Scotia) under stones along the banks of the Salmon river.
The larvae left the water about the last of July or the first of
August. They constructed their pupal chambers in damp places
under stones. Matheson found many pupae and larvae in such
situations about August 7, and succeeded in rearing one adult from
these pupae. It emerged on August 10. From this information it
seeems probable that the pupal state may last two weeks or less.
In July and August I have found adults and larvae of all stages in
lakes and streams. Larvae kept under laboratory conditions for a
period of four to six weeks have not been seen to moult. The
data are insufficient to warrant a guess as to the probable length of
the life cycle, but they suggest that the complete development may
occupy a period of more than one year.

Recently Hinton (1936) lias made some observations on the
biology of Dryops luridus Er., a European species. Eggs were de-
posited from May to July. Some of the eggs were found inserted in

644 The University Science Bulletin

fresh and partly decayed stems and leaves of the water mint,
Mentha, and still others were found at the bottom of the rearing
jar. Hinton considered that oviposition normally occurs in plant
tissue, but that the overcrowded conditions forced some females to
lay their eggs free in the water. The eggs required about fifteen
days to hatch. Rupturing of the chorion took place by body move-
ments. According to Hinton (p. 72), "Two or more days before
eclosion the larva may be seen frequently to shift to a slight ex-
tent its position inside the egg. It often actively moves its legs
and mouth parts. No special apparatus exists for breaking the
chorion, as has been observed for other beetles, such as Dytiscus.
Probably no fluid is secreted, for if the egg be marked it is found
to break open at places where the head of the larva has not been.
On emerging, the larva usually begins to tunnel in the partly de-
cayed stem or leaf in which the egg was laid. The larvae, both in
nature and in captivity, seem to prefer partly decayed plant tissue.
The duration of the life cycle and the number of instars has not
been determined, nor have I been able to obtain pupae. I believe
a complete life cycle probably requires two years. West (1929, p.
18) points out that in all probability the life cycle of Psephenus
lecontei Lee. requires two years in northern latitudes."

In a related species {Dryops auriculatus Geoffr.), Brocher (1913,
p. 227) has noticed the females depositing their eggs in damp places
out of the water, though this may not be the usual habitat for the
species.

Respiration and Habits of the Larva

The larvae of the various species of Stenelmis which I have ex-
amined are entirely aquatic and breathe by means of caudal fila-
ments. The respiratory organ consists of three tufts of filaments
which, in later instars, arises from a common stalk and which may
be protruded from the caudal opening of the abdomen. The most
important contribution to this subject is by Susskind (1936). This
author treated the morphology and function of the respiratory
system of several instars of Stenelmis quadrimaculata Horn. {Stenel-
mis sulcata Blatch.) She observed that the larvae expand and con-
tract the filaments rhythmically when they are subjected to ab-
normal conditions such as rapid increase in temperature of water,
exposure to light, high carbon dioxide tension, or low oxygen ten-
sion. The rhythm of expansion might be slow or rapid, depending
upon the various conditions to which the larvae were subjected.
The larvae often gathered themselves in a mass and remained quiet
in this position for long periods. At such times the filaments usually

Sanderson: Species of Stenelmis 645

were retracted, and she suggested that under these circumstances
respiration might go on without their use.

Adaptation of the larva to gradually changed conditions is gener-
ally effected without unusual disturbance to itself. If, however, the
larva is suddenly subjected to warm water, the body is rapidly con-
torted in much the same manner as a mosquito larva.

The paper by Miss Susskind presented the various methods by
which the tracheae of the larvae were prepared and studied. Further-
more, she has traced the various branches to the different parts of
the body. The average number of filaments given for the middle
tuft of caudal filaments is 55, and for the lateral tufts, 35. Two
tracheoles, twisted about each other, are in each of the filaments.
According to this author the spiracles do not appear until the last
larval instar. There are ten pairs, two in the thorax and eight in
the abdomen. The position of the spiracles is ventrolateral with
one pair each in the mesothorax and metathorax, and one pair in
each of the first eight abdominal segments. The aperture of the
spiracle is described as being closed by two lightly sclerotized mem-
branes which meet in a straight line parallel to the long axis of the
body. The internal edges of the membranes are fused. She thought
that the membranes prevent the entrance of water into the tubes
and probably act as diffusion membranes.

By way of summarizing her paper, Miss Susskind has given the
following tabular arrangement:

1. Both larvae and adults of Stenelmis sulcatiis {quadrimaculata Horn)
occur in marl concretions and are permanently submerged, never coming to
the surface for air.

2. The larval tracheal system has fewer branches in the first instar than
in all older larvae.

3. In the first larval instar neither spiracles nor spiracular tracheae are
present.

4. The intermediate instar studied has spiracular tracheae, which are about
half developed.

5. Last instar larvae have completely devolped spiracular tracheae and
ten pairs of spiracles. The spiracular apertures are permanently closed by
weakly chitinized membranes.

6. Larvae of all instars possess three tufts of caudal filaments at the
posterior end of the main tracheal trunks. These tufts are protrusible through
the tip of the last abdominal segment.

7. Intermediate and last instar larvae possess a caudal chamber associated
with the caudal filaments.

8. The tracheal system of all larval instars is a closed one.

Hinton (1936) has figured a ventral view of the tracheal system
of the larva of Dryops luridus Er. which, however, belongs to a

646 The University Science Bulletin

different subfamily from Stenelmis. This constitutes practically
all that is known regarding respiration and respiratory systems of
the larvae of this family.

I have observed in the field and laboratory the larvae of Stenelmis
quadrimaculata Horn in marl concretions. They occur in immense
numbers in Black Lake (Cheboygan county, Michigan), where
they may be taken from tlieir burrows in the concretions. Some-
times at the ends of these burrows a mass of larvae, often of various
stages, is found. The larvae of thq stream-dwelling species of
Stenelmis crawl about beneath stones, in sand and in debris lodged
in the stream. Hinton (1936) records the larvae of Dry ops luridus
Er. as having been taken, on one occasion, from damp places some
distance from water. He suggested that since the larvae were in
their last instar, the entire life cycle might take place out of water.
I have collected larvae from running waters of all types, and also
from the waters of lakes. Hinton's records are, therefore, interest-
ing, and unusual if we consider tlie preferences of the great bulk of
our Dryopidae.

Respiration and Habits of the Adult

Texts and other papers which discuss the respiration of aquatic
insects rarely mention that it is unnecessary for adult Dryopids to
come to the surface of the water to breathe. Except for the genus
Haemonia (Chrysomelidae), other Coleoptera, although they may
descend beneath the surface of the water to deposit their eggs (ex.
Psephenidae), or may use this medium as a natural habitat (ex.
Dytiscidae), are not known to remain there for indefinite periods
without returning to the surface for air.

The body of a submerged dryopid is surrounded by a film of air
which is held to it by the hydrofuge hairs. This silvery region is
termed the "plastron" by Brocher, Muttkowski and others. This
arrangement is not alone responsible for the insect's ability to re-
main under water indefinitely, for other aquatic Coleoptera have
similar air-carrying capacities, yet they can remain under water
only temporarily.

The researches of Brocher (1912) have contributed much regard-
ing the respiration of adult Dryopidae. I shall make no attempt
to cover his extensive paper on this subject, but hope to present
some of his interesting experiments and conclusions, especially re-
garding Stenelmis canaliculata (GylL).

Brocher, at first, studied certain European species, including the

Sanderson: Species of Stenelmis 647

above Stenelmis, under laboratory conditions. The jars in which
they were placed contained sand, stones with calcareous incrusta-
tions, and some aquatic plants such as Hypnum. They were kept
in a cool, well-lighted place, but out of the direct rays of the sun.
Under these conditions, the plants liberated a great deal of oxygen,
the water temperature did not rise, and there w^as an abundance of
food. The beetles behaved as they did in their natural habitat and
remained in perfect health. Some of the adults were placed in a
small jar, the mouth of the jar was closed with muslin and sub-
merged in a larger jar of water. They remained thus for 140 days,
when the experiment was interrupted, at which time they were still
in perfect health. During this time the adults were not always ob-
served with a bubble of air at the tip of the body, nor were the
beetles found floating inactive at the surface of the water. When
the conditions of existence became unfavorable, such as too few
plants in the jar, or too feeble illumination, the beetles were dis-
turbed. Some moved about with a bubble at the tip of the ab-
domen; they were often seen rising to the surface, supported here
by the bubble. If the bubble burst, the insect immediately fell to
the bottom of the aquarium. Brocher concluded, after this obser-
vation, that the beetle did not come to the surface for air. He
mentions that these insects are the first to die when the water of
the aquarium is insufficiently aerated.

Proof that Dryopidae obtain some of the oxygen necessary for
respiration from plants presented itself to Brocher on two occasions.
Adults were observed, upon encountering a bubble of air adhering
to the plant, "to eat with frenzy at this spot." The silvery region
of the head absorbed some of the oxygen, gradually relaying it to
other parts of the body. He did not observe air in the mouth of
Stenelmis, which led him to suppose that the absorption of oxygen
must have taken place in other regions of the body.

He further observed that the Dryopidae were able, while feeding
on plants, to gather quantities of oxygen which oozed out of the
tissues. The oxygen, apparently, was absorbed by the silvery sur-
face of the body. The fact was mentioned that it was important
to the insect not to take up too large a quantity of gas at one time
since the specific weight of its body would be lowered. In this event
the beetle might float to the surface of the water once it lost its
hold ui)on a support.

After having studied the habits of healthy Dryopids, Brocher
thought it worth while to study certain beetles in which the functions

648 The University Science Bulletin

of respiration had been disturbed. One experiment was to force the
beetles to stay in a poor oxygen medium, such as boiled water, or
in water charged with carbonic acid. When the bubble of air ap-
peared at the end of the body, it was removed. In addition, the
silvery region of the body was mechanically removed or the end
of an elytron w^as cut off. When they were transferred back to the
proper conditions of the aquarium, the insects appeared to be more
active than those that were kept in the normal state.

It was found that when the hairs which support the silvery region
of the prosternum, mesosternum, and metasternum were completely
removed by scraping with the point of a needle, the beetle died
within one or two days. When only the silvery region of the pro-
sternum was destroyed, the beetle was able to survive for a longer
time and apparently in good condition. The facts, according to
Brocher, appear to demonstrate that the silvery regions fulfill an
important physiological function which can only be respiratory.

The respiratory system is described by Brocher as being composed
of the silvery hydrofuge surface, the dorsal space under the elytra,
and the tracheal system. In Stenelmis the silvery region w^as de-
scribed as occupying the dorsal surface of the head and prothorax
and all of the ventral face of the body, including the femora. I
have also observed this region covering the tibiae. When these
surfaces were examined under the microscope, many granules w^re
found, each provided with a long hair. Brocher regarded these
granules as organs destined to capture the minute bubbles of gas
which by chance come into contact with the silvery regions. The
bubbles came to rest collectively on the granules and little by little
were absorbed.

Brocher described nine pairs of spiracles in the adult. The first
pair was situated in the membrane between the prothorax and the
mesothorax. These were termed the "mesothoracic spiracles." The
"metathoracic spiracles" were situated in the lateral membrane of
the metathorax opposite the coxal cavity. They are much less
prominent than the mesothoracic ones. Raising of the elytra and
wings revealed seven pairs on the dorsal face of the abdomen.

The hydrofuge hairs are continuous, from the ventral surface of
the abdomen, along the groove under the edge of the elytra, and to
the spiracles. By this route the oxygen may pass directly to the
spiracles where it can be used for respiration or to the subelytral
space for storage. It is thought that the oxygen which comes to the
silvery regions of the prothorax, by way of the mouthparts, arrives

Sanderson: Species of Stenelmis 649

at the mesothoracic spiracles which may act as inspiring spiracles.
Frequently, however, they may serve for expiration, for bubbles
have been observed to issue from them. The metathoracic spiracles
have a direct relationship with the hydrofuge regions or the ventral
surface of the body, as has been pointed out for the abdominal ones.

Frequently Brocher noticed that when a Dryopid was placed on
its back and further disturbed by teasing, a bubble of air appeared
on each side of the posterior coxae. He was led to believe, from this
observation, that expiration also took place in the abdominal spir-
acles. By pressing lightly on the abdomen, I have noticed that a
bubble of air appeared at the tip of the body. Another specimen,
when turned on its back, released two bubbles of air from the
region of the left hind coxa, and another from the left middle one.

I have observed the adults of Stenelmis quadrimaculata Horn
feeding upon algae in marl incrustations. Specimens taken from a
stream and brought into the laboratory with dead leaves were found
to "graze" over both surfaces. They were, apparently, feeding upon
an algaelike gro^i;h. When no leaves or stones containing their
food were placed with them, they proceeded to feed upon the debris
on one another.

The tarsi of the adult Stenelmis are terminated by two long claws
which enable them to cling to stones in the swiftest of streams.
They seldom come to the surface during the day, but at night the
winged species often leave the stream, pond, lake, or other natural
habitat and take wing. They have been taken at lights by hundreds
in various localities. Frequently several species may be observed
in these flights. This indicates a positive phototropism, though dur-
ing the day they generally prefer the dark recesses of their habitat.
In at least one instance, however, I have found adults of a Dryopid,
Heterlimnius corpulentus (Lee), during a bright day, resting on
stones jutting out of a stream.

Description of the Larva

Matheson (1914) briefly characterized the larva of Stenelmis
bicannata Lee. The full-grown larva is described as having the
head and terminal segment of the abdomen nearly black. The
antennae are briefly described and measurements are given for each
segment. Following Matheson, we have the next important con-
tribution, on larvae of Dryopids, by West (1929). This author dis-
cusses eight types of larvae and, in addition, the larva of Psephenus
herricki (DeKay) (Psephenidae) . His description of an unidentified
Stenelmis follows:

650 The University Science Bulletin

"Larva elongate, of the form of an attenuated hemicylinder, /. p., with
ventral surface nearly flat, but with dorsum arched, so as to appear in cross
section semicircular. Median dorsal line but faintly indicated; not elevated
into a carina of any sort. Prothorax one and one-half times the length of the
succeeding segment. Seen in dorsal view, margins of all body segments except
the last appear bisymmetrically rounded, the margin of the prothorax alone
being very slightly sinuate. Posterior abdominal segment subconical ; the
dorsal sclerite rather abruptly narrowed posterioi'ly and terminating in two
sharp points. Ventral sclerite occupying posterior hall' of segment, ol the
form of a cjuadrangle with an equilateral triangle attached posteriorly; with
two elongate, slender appendages, bearing numerous sjiines, and recurved at
the tips. Entire doisal surface of the larva, except the head, covered with
a close sprinkling of fine tubercles, each of which tends to give rise to a
minute, backward directed spine. These tubercles tend to be replaced by
spines on jiosterior segments. Those spines along the posterior margin of each
segment arc more stout and elongate, forming a ciliate border to the segment.
Larva slightly broadest in the region of the first two or three abdominal seg-
ments; diminishing slightly in size both anteriorly and posteriorly from this
point.

"Head large, exserted, rather consi)icuous, being nearly three fourths as
broad as prothorax; with conspicuous Y-shaped epicranial suture visible
dorsally. Antennae short, composed of three segments; the first rather deeply
inserted and but slightly longer than bioad, the second much more slender,
fully twice the length of the first and, seen laterally, more expanded at distal
end than at the l)ase; the third duplicate, being composed of two small,
transjiarent articles arranged side by side, the one gradually tajiering, the
other more robust and of the same diameter throughout. No terminal spine
has been demonstrated on either of these articles. Ocelli situated behind the
antennae, each group composed of apparently fi\'e units. Labruni simple,
with lateral margins evenly rounded; slightl.v emarginate anteriorly. Anterior
and lateral margins, as well as entire upi)er surface, beset with rather stout
spines. Mandibles subtriangular, distinctly tridentate, and bearing a large
"taste-brush" (prostheca) on the inner surface at about one third the distance
from base to tip. Outer surface bearing at least two branching spines. Max-
illae small, with four-segmented palpi. Stipes, near base of palpus, bearing
at least one stout branching spine. Small area on distal portion of last
palpal segment ])rovide(l with i)a]iillae, evidently tactile in function. Lacinia
lying close upon galea and of the usual form, i. c, resembling a truncate
sleeve, or cylinder, open along the exposed surface and adorned distally with
a variable mmiber of stout spines. Galea without characteristic form, bear-
ing distally several stout curving spines. Labium membranous, nearly twice
as long as broad, and bearing three-segmented palpi, of which the third
segment is much the smallest and bears distally a small sensory area, where
are located usually four tactile iiapillae. Outside of, and opposite to first
palpal segment may be found, on each side, a short, studded spine. Mentum
rather clearly bounded by a suture at the base of the palpi. Distal portion
of labium evenly convex and adorned with a great number of fine, tactile
hairs.

"Legs composed of the usual parts and adorned with a rather scanty cover-
ing of spines. Coxae somewhat excavated anteriorly, and thus adapted for

Sanderson: Species of Stenelmis 651

reception of femora. Distal portion of excavation cushionlike in appearance,
with a mass of small, closely ranged tubercles. Claw bearing on its inner sur-
face a slender and sometimes curving spine. Gills caudal, arranged in three
principal tufts and capable of extrusion between the sclerites of the pcsterior
abdominal segments. Spiracles situated laterally, one pair on the mesothorax
and one pair on each of the first eight abdominal segments. Length of mature
larva, 6.5 mm. Greatest width (at first abdominal segment), .91 mm.

"Description drawn from several specimens taken at Walnut Creek, Michi-
gan, and a.'ssigncd to Stenelmis since sevei'al adults of that genus were associ-
ated with them. Also the larva of Sfenchnif; bicarittatus Lee, as figured by
Matheson ("14), comes closer to tliis type than to any other studied so far."

Plate I, figures 8 and 9, and Plate IV in West's paper illustrate
larvae and certain morphological characters. In West's discussion
of the larvae, a comparison is made of the larval types and he at-
tempts to correlate structures of the larvae with their habitats.
Comparisons are made chiefly with Psephenus, Helichus, Elmis
(European) and Stenelmis.

The lateral lobes of the body segments of Psephenus are developed
to a high degree, but, when one proceeds through the series to
Stenelmis, the lobes are found here to be entirely lacking. Com-
parison of the respiratory apparatus, especially position, is of some
significance in the series. In Psephenus the respiratory gills are
ventral; in all others they are caudal. The structure of the caudal
abdominal segment is described by West as being strikingly similar
in most of the larvae having caudal gills. Position of the head in
Psephenus and Helichus is beneath the expansion of the prothorax,
but in others the head is visible from the dorsum.

Following this discussion West attempts to show natural group-
ings within the family on the basis of larval characters. An in-
terpretation is given from an evolutionary point of view. A key, in-
cluding several European forms, is presented for the few larval
types.

Boving and Craighead (1930), in their extensive paper on the
principal larval forms of Coleoptera, figure a mandible, head and
antenna, maxilla, and a lateral view of the ninth tergite and tenth
sternite of Stenelmis crcriata (Say). Descriptions of the parts are
not included.

Finally, Bertrand (1935) briefly describes some characters of four
larvae, thought to belon'g to Stenelmis, from Java. Only the antenna
and mandibles are figured.

Comparative studies of the larvae, within a genus of Dryopidae,
have not been made. The larvae of some genera still are unknown,
but, when most of the larval types have been discovered and studied.

652 The University Science Bulletin

we may have a better understanding of the phylogeny of the entire
family.

Description of the Pupa

Matheson (1914) gave us our only figure and description of the
pupa of Stenelmis. According to this author:

"The pupa {Stenelmis bicarinata Lee.) is soft, white in colour, the thoracic
segments, wing pads and legs being slightly dark in colour. This darkness
deepens with the age of the pupa. Length, 3.4 mm.; width at base of wing
pads, L2 mm.

"The head lies incurved under the prothorax, the developing mouth parts
showing very distinctly. The eyes are small and almost black. The antennae
lie in front of the eyes and extend under the pronotum, thus being concealed
from the dorsal view. The wing pads are prominent.

"The pronotum is large and rather densely covered with fine colourless
setae. The outer angles of the anterior margin each bear a long, curving spine,
measuring .32 mm. The posterior outer angles, also, each bear a spine measur-
ing .28 mm. These spines are yellowish-brown in colour. They support the
pupa in its chamber, preventing it from coming in contact with the coarse
grains of sand.

"The abdomen is composed of nine segments, the posterior margins on the
dorsal side being strongly elevated into narrow ridges. These ridges are
clothed with numerous short setae. The ninth segment bears on its dorsal
side two curving, stout cerci, measuring .4 mm. in length. The cerci almost
completely conceal the segment from which they arise. The pupa rests on
its back in the pupal chamber. By the aid of the prothoracic spines and cerci
together with the numerous short setae arising from the raised ridges of the
abdomen, injury from the roughened walls of the pupal chamber is avoided.
The posterior margins of the wing pads and the legs are also provided with
many short setae."

GEOGRAPHICAL DISTRIBUTION

The family Dryopidae is not a large one, there being some 600
species representing approximately 80 genera known from the en-
tire world. Their distribution extends to all of the continents, and
nearly every locality, through the tropical and temperate zones, is
productive of from one to many species. The greater number of the
known species appear to come from the Oriental region, and from
the East Indian group of islands. In a number of instances a genus
may be confined to a definite region. The genera Pachelmis Fairm.,
Helminthocharis Grouv., Lophelmis Fairm, and a number of others
are recorded only from Africa and Madagascar. Phanocerus Sharp
is known only from North and South America and the West Indies.
However, other genera are more widely distributed and not con-
fined to a limited region. Dryops Oliv. is found in Europe, Asia,
North and South America, and Africa. Thus, it might be said to be
found on all continents with the possible exception of Australia.

Sanderson: Species of Stenelmis 653

The genus Stenelmis Duf. is one of the larger genera of the
family, sixty-nine species having been described. Nearly one half
of the species are from the Oriental region and the East Indian
Archipelago. Sumatra, alone, has ten species. The United States
is second in importance to the Orient with one third of the species.
The rest occur over wide areas, three recorded from Europe, ten
from Africa, and one each from Damascus, French Guiana, Borneo,
and Costa Rica. It is of interest to note that one species, S. minuta
Grouv., has been recorded both from Sumatra and Java.

Although there are, no doubt, many more new species to be dis-
covered from one region or another, this distribution suggests that
the genus may have arisen in the Orient.

In the United States the species of Stenelmis, with one exception,
are distributed east of the 100th meridian from Canada south to
Florida and Texas. S. crenata (Say) is distributed over the whole
of the eastern United States from Quebec to Texas. On the other
hand, S. sexlineata Sand, is, apparently, a middle western form, hav-
ing been found only in Kansas and Texas. Some of the species
occur in large numbers in small areas with an occasional specimen
found in other localities. This is especially true of S. lateralis
Sand., which is abundant in the streams of the Ozark Mountain
region. A few specimens have been taken in Pennsylvania; Virginia;
Tennessee; and Ireland and Lucedale, Miss.

Until more is known of the biology of this genus, it would be out
of place to generalize regarding the habitat preferences of the vari-
ous species. However, some of the species seem to prefer clear and
cold streams; others muddy and warm ones. Other species are
found in both. Two species, S. quadrimaculata Horn and S. doug-
lasensis Sand., have been found in lake water and, from available
information, they seem to be confined to this habitat.

In the great expanse between the 100th meridian and California,
no species of this genus has been recorded, although considerable
collecting has been done. However, S. nubifera Fall, our only
western species, is distributed along the Pacific coast from Southern
California to Oregon.

GEOLOGICAL DISTRIBUTION

A very few fossil Df'yopidae or Psephenidae have thus far been
recorded. Apparently the first fossil Dryopid to be made known
was Dry ops (Parnus) prolifericornis Fab. from the Pleistocene of
Galicia. This species, identified in 1894, was associated with the
recent species D. prolifericornis Fab. of Europe, which is now rec-

654 The University Science Bulletin

ognized as a synonym of D. auriculatus Geoffr. In 1911 Wickham
recorded Dry ops eruptus (now assigned to the genus Helichus) from
the Florisant. The following year, he named Dry ops tcnuior (also
placed with Helichus), and named and described the genus and
species, Lutrochites lecontei, both species of which came from the
Florisant. The first Psephenid, made known in 1911, was Psephenus
lutulentus Scudder, described from the Florisant of Colorado. Con-
cerning this species, Scudder says, "The species described below,
from the Oligocene of Colorado, is the only extinct form of this
family yet known." The Florisant of the Miocene is the correct
horizon, no doubt, rather than the Oligocene. This constitutes
virtually all that has been made known of fossil Dryopidae and
Psephenidae.

BIBLIOGRAPHY

1894. Parnus prolijerlcornis Fab., Lomnicki, Miis. Dziedusz. IV, 74, t. 6 f. 57.

1900. Psephenus. lutulenius Scudder. Mon. U. S. Geol. Survey XL, p. 94.

1911. Dryops eruptus Wickham. Bull. Amer. Mus. 30, p. 56.

1912. Dryops tenuior Wickham. Bull. Nat. Hist. S. U. I. VI. p. 16.
1912. Lutrochites lecontei Wickham. Bull. Nat. Hist. S. U. I. VI, p. 16.

Stenelmis Dufour (1835)

1835. Dufour, Ann. des Sci. Nat. Second Series Tome III, p. 158.

1847. Erichson, Naturg. Ins. Deutsch. Ill, p. 534.

1851. Sturm, Deutschl. Ins. XXII. p. 30.

1852. LeConte, Proc. Acad. N. Sci. Phik. VI, p. 44.
1859. Motschulsky, Etudes Entom. VIII, p. 50.
1870. Hoin, Trans. Amer. Ent. Soc. Ill, p. 39.
1872. MuLsant et Rev, Hist. Nat. Col. Fr. V, p. 128.
1904. Ganglbauer, Kaf. Mitteleur. IV, I, p. 107.

1907. Reitter, Zeitschr. p. 483.

1908. Zaitzev. Horae Soc. Ent. Ross. XXXVIII, p. 299.
1910. Zaitzev, Junk Col. Cat., par. 17, p. 21.

1910. Blatchley, Col. Indiana, p. 680.

Genotype. Elmis canaliculata Gyll., 1808.

DESCRIPTION OF THE GENUS STENELMIS

Elongate, convex or subdepressed species. Head retractile and
protected beneath by the prosternal lobe. Antennae eleven-seg-
mented, the segments gradually broader toward the distal one, the
basal one or two wider and more rounded than succeeding ones.
Maxillary palpi four-segmented, the labial palpi three-segmented.
Mandibles bidentate at apex. Eyes denuded. Pronotum usually
quadrate or longer than broad, convex, with a median longitudinal
sulcus, and on either side a rounded or elongate basal tubercle. A

Sanderson: Species of Stenelmis 655

tubercle in front of tlie basal one generally rounded or elongate,
though occasionally indistinct anteriorly from the general surface of
pronotum. Apical angles of pronotum prominent. Elytra punctate-
striate with the third interval usually elevated at base, the sixth
elevated and usually carinate for nearly its entire length. Front
coxae rounded, without trochantin, widely separated by posterior
lobe of prosternum which fits against a groove in the mesosternum.
Middle coxae rounded, more widely separated than the prosternal
coxae. Posterior coxae transverse, dilated internally, with their
separation less than that of the middle coxae. Ventral abdominal
segments five, the apical one with its apex more or less emarginate.
Legs slender, the hind legs longer than either the middle or anterior
ones. Tarsi five-segmented, the last segment generally longer, but
never more than slightly shorter than the four preceding segments
combined. In our species, anterior tibiae without brush of to-
mentum on inside of apical half.

Secondary Sexual Characters of Stenelmis

In all of our species the males have been found to possess a
spinous ridge or a row of spinules on the inside of the middle tibia.
The spines are variable in number within the species. They usually
extend posteriorly for a short distance from near the middle of the
tibia. The females have the inside of the middle tibia smooth.
Both middle and hind tibiae of Stenelmis nubifera Fall have been
found to possess the same type of spination. Another character,
of sexual importance, which has been found in at least one species,
Stenelmis quadrimaculata Horn, is a white labro-clypeal band.
This membranous band is entirely absent in the males of the species,
but very distinct in the females.

Specific Characters in Stenelmis

In this genus, in which most of the species are closely related, the
search for characters l)y which to segregate species has been a
rather difficult task. Since no key has been published to include
all of the known members of our fauna, a means was first sought
to separate the species into groups. Inasmuch as all of our species,
with one exception, are confined to the eastern half of the United
States, a study was made of the Pacific Coast Stenelmis nubifera
Fall to see if it presented any marked differences from those of
the East. Examination shows that the first elytral stria of Stenel-
mis nubifera Fall is incomplete, and does not extend to the apex
of the elytron. Furthermore, the granulations of the head, pro-

656 The University Science Bulletin

notum, and legs are conspicuously elongate, contrasted with the
rounded granules of the other species. Other characters were also
noted, such as the type of maculation, etc. One species was thereby
eliminated, and study was then made of the remaining ones to see
if they presented any group characters. An examination showed
that they could be differentiated into two fairly well-defined groups,
based upon the comparative length of the last tarsal segment with
the four preceding segments combined.

The groups are accordingly designated as Nubifera group with
one species; Crenata group with eleven species; and the Sinuata-
humerosa group with fifteen species. The members of the Crenata
group have the last tarsal segment no longer than the four pre-
ceding segments combined. Those of the Sinuata-humerosa group
have the last segment distinctly longer than the combined length
of the preceding four. The longer last segment invariably has the
claws more robust.

Descriptions of the different species have been arranged, in this
paper, in a logical manner and the scheme has been consistently fol-
lowed throughout. After a species has been traced to its position
in the key, it should be carefully compared with its description.

The following arrangement has been used in all descriptions:

Size. The total length of the species, indicated under size, may
fall short of the total length of the pronotum plus elytra. This is
explained by the fact that the pronotum is not in the same plane
with the elytra, but is slightly deflexed. Measurements of the width
are from the greatest width across the elytra.

Form and color. The form of Stenelmis may vary somewhat from
one species to the next, but in degrees that make description difficult.
In general, however, the range is from nearly subdepressed to con-
vex. The body is usually slender, and with the sides of the elytra
nearly or quite parallel.

Head. All of our species, except S. nubifera Fall, have a dark
median longitudinal band on the head with two lighter bands, nearly
of equal width, on the inside of the eyes. The bands may be in-
distinct if the specimens have the head covered with earthy matter.
In several cases the color of the antennae and palpi aid in segre-
gating certain species. The length of the antenna is helpful in
certain groups. Several of the last antennal segments in many
specimens are clothed with very fine, yellow hairs and a few stiff
ones near the apices of these segments. This character has not
been found of specific value, for the hairs may be present or ab-
sent within a series of a single species from one locality.

Sanderson: Species of Stenelmis 657

Pronotum. There is every gradation in tlic prunotum from one
in which the median longitudinal sulcus or groove is very deep,
with the sides nearly straight, to one in which the groove is barely
evident. There is, likewise, much variation in the size and shape
of the tubercles at base of pronotum between posterior angles and
sulcus. In some species this tubercle is carinate posteriorly; in
others the carina is obsolete. The tubercle immediately preceding
the basal one is designated as the anterior tubercle and is variable
to some degree. It is rounded and may be very prominent to in-
conspicuous. The oblic]ue transverse impression separates the lat-
eral tubercles in a varying degree and is occasionally deep immedi-
ately on the inside of the basal tubercle. Behind the anterior angle
of the pronotum is a small, transverse ridge, continuous with the
lateral margin of pronotum, and variable in its distinctness. In two
species, S. tarsalis n. sp. and »S. concinna n. sp., the anterior tubercle
barely unites with this transverse elevation. In some species the
crenation of the lateral margin of pronotum is better marked than
in others, although this character seldom aids in the separation of
closely related species. The pronotum, as well as the elytra, is
covered with fine, usually close, and decumbent hairs. These appear
to be more distinct on the margins of the median sulcus and the
base of the third elytral interval. The pronotum is nearly always
longer than wide, and especially so in the Sinuata-humerosa group.
In S. tarsalis n. sp., however, the width is slightly greater than the
length.

Elytra. The keys, to a great extent, are based upon the macula-
tion of the elytra. Therefore, it is necessary to start with a speci-
men in clean condition, or nearly so. However, two species are
known in which the elytra are entirely without spots or vittae. One
species is known by the presence of six longitudinal vittae on the
elytra. Other divisions within the two major groups have the hu-
meral spot covering the umbone of the elytron, or the spot may be
confined to the inside of the sixth elytral interval. When the vitta
is divided into two spots, the basal one is termed the basal spot, the
other is the apical spot. Rarely do individuals of the same species
show every gradation between an entire vitta and one in which the
spots are separate.

The first interval is designated, in this paper, as the one between
the elytral suture and the first stria. In Stenelmis nubifera Fall the
first stria is short, with the fourth interval elevated at base. The
seventh interval is also elevated in this species. In all others of the

42-4141

658 The University Science Bulletin

genus in our fauna the first stria continues to the apex of the elytron.
The third interval is elevated at base in all but one or two of the
remainder of our species. The sixth interval is always more or less
carinate from the base for about five sixths of its length.

Venter. The apex of the last abdominal segment is always emar-
ginate, although the emargination may be partially covered above
by a thinner part of the segment. In aS. nubifera Fall the width of
this emargination is about twice that of the greatest width of the
last tarsal segment. In all others the width may be less than, to
but little more than, that of the tarsal segments. The value of this
character is especially significant in such closely related species as
S. taTsalis n. sp. and >S. concinna n. sp. The color of the venter is
variable and is of little importance. In perfectly clean specimens
the venter is a beautiful irridescent blue.

Legs. The character of the last tarsal segment is of fundamental
importance in the separation of our groups of Stenelmis. The Sinu-
ata-humerosa group, in which the last segment is longer than the
preceding four combined, has this segment more suddenly enlarged
distally than in the Crenata group. The claws in the latter group
are slender and much less robust than in the other group. In two
species the lower margin of the last tarsal segment is triangularly
produced. This margin in all other species is straight or but little
produced. The color of the tibiae and tarsi is useful in separating
some species. With exception of *S. douglasensis n. sp. and S. gj-ossa
n. sp., all of the species in our fauna have the pronotum, venter, and
legs granulate. The granulation of the femora refers to the larger
ones on the outer surface. These are frequently intermixed with
many fine granules. The hind tibiae are always longer than the
middle and anterior ones. The anterior tibiae are longer or equal
to the middle tibiae.

The Male Genitalia of Stenelmis

The genitalia of the males of Drypos, Helichus, Elmis, and a few
other genera have been used to some extent in classification, but to
the writer's knowledge, the genital structures of but one Stenelmis
{nevermanni Hntn. 1935) have been figured and described for this
purpose. Since some of the species of Stenelmis are difficult to
segregate by external characters, a study of the male genitalia
shows that this organ is of some importance. The genitalia are
quite simple, consisting primarily of a large cylindrical basal piece,
a median, and two lateral lobes. The bases of the lobes are inserted

Sanderson: Species of Stenelmis 659

for a short distance inside the posterior opening of the basal piece.
The median lobe is longer than the lateral ones.

In our species of Stenelmis four general types of genitalia are
represented: (1) lateral lobes of aedeagus without internal sinua-
tion, (2) lateral lobes sinuate internally, median lobe without lateral
processes, (3) lateral lobes sinuate, median lobe with lateral proc-
esses evenly rounded, and (4) lateral lobes sinuate, median lobe
with lateral processes subangulate anteriorly. The first type is
found in S. nubifera Fall and the remaining types are found among
the species in the Eastern United States. It must be pointed out
that the genitalia are not to be depended upon entirely for the
identification of species, but should be used in conjunction with the
keys and descriptions. For example, one type might be exhibited
in each of our chief groups of Stenelmis and the resemblance be-
tween the genitalia of two species, each in a separate group, may
seem close.

Technique for Dissecting Male Genitalia

The technique for extracting the genitalia is simple and is as
follows: The male is determined by the presence of the small spines
on middle tibiae, all labels are removed from the pin, and the speci-
men is immersed for several minutes, or as long as necessary, in a
5-10 percent hot alcohol solution. After the beetle is sufficiently
relaxed, it is held between the thumb and forefinger; a small, curved
dissecting needle is inserted between the last dorsal and ventral
abdominal segments; and the genitalia and associated parts are re-
moved. A few minutes' boiling in a weak solution (5-10 percent) of
caustic potash, then washing in distilled water, will clear the struc-
tures sufficiently for study. It has been found very convenient to
mount the genitalia below the dissected specimen upon a small
celluloid slip with the genital parts immersed in a drop of com-
mercial diaphane (euparol). Thus the genitalia is always with
the specimen and is easily removed by dissolving the diaphane in
ninety-five percent alcohol. The genitalia may also be mounted on
glass slides in Canada balsam or diaphane and covered with a cover-
slip. Another method is to put the genitalia in a drop of glycerine
in a small vial which is of the size that it may be kept on the same
pin with the dissected specimen. The pin is passed through the cork
of the vial.

660 The University Science Bulletin

SYSTEMATIC TREATMENT
Key to Genera of Dryopidae of the United States

At the time of this writing, the world genera are in the process of
being revised by Mr. Howard E. Hinton, Berkeley, Cal., after which
this key may be subject to modification. No attempt is made to
extend it beyond the limits of the United States, and for that reason
it is applicable only to the forms within this region. It is given
simply to show the relationship of Stenelmis to the other genera.

At present it is considered that the Dryopidae are divided into
three subfamilies, as follows:

SUBFAMILIES OF DRYOPIDAE

1. Head not entirely retractile; anterior lobe of presternum ill defined or absent; mouth-
parts exposed; posterior coxae dilated and partly protecting femora Larinae,

Head usually retractile, usually protected beneath by prosternal lobe;n:outh parts gen-
erally concealed 2

2. Anterior coxae, with transverse trochantin Dryopinae,

Anterior coxae globular, without trochantin Elminae,

Lminae

Antennae clavate Phanocerus,

Antennae not clavate Lara,

Dryopirme*

1. Body rounded; last segment of maxillary palpi hatchet-shaped Lutrochus,

Body elongate, oval; last segment of maxillary palpi slender 2

2. Second segment of antennae produced into an earlike process 3

Second antennal segment not thus produced 4

3. Antennae approximate; thorax with a deep-cut, sharp-edged, longitudinal line on each

side Dryops,

Antennae widely separated; thorax not as above Helichus,

4. Antennae distant Pelonomus,

Antennae approximate Throscinus,

Elminae

1. Anterior tibiae without a patch of tomentum Stenelmis,

Anterior tibiae with tomentum 2

2. Antennae of less than 10 segments 3

Antennae of 10 or 11 segments 4

3. Antennae 7-segmented Macronychus,

Antennae 8-segmented Zaitzevia,

4. Maxillary palpi 3-segmented 5

Maxillary palpi 4-segmented 6

5. Sublateral carinae present Elmis,

Sublateral carinae absent Narpus,^

* The genus Oberonus Csy. has been omitted from the key to the Dryopinae. At the
present time I am not of the opinion that it should be made distinct from Pelonomus until
additional specimens are examined to verify the characters set forth by Casey. I have
examined the type, and it is true that the structure of the intermediate coxae is radically
different from Pelonomus. However, this may be no more than an imperfectly developed
structure. The type of Oberonus is a male and the genitalia are exactly the same type as
Pelonomus.

tThis genus was incorrectly placed with the Dryopinae until its correct position was
established by Hinton (Ent. Mo. Mag. Vol. 72, 1936, p. 57). I have examined the im-
perfect type of this species, and in addition, several perfect specimens. The type of antennae,
as well as the globular anterior coxae, unmistakably place it in the Elminae.

Sanderson : Species of Stenelmis 661

6. Head free, prosternum not lobed in front Ancyrnnyx,

Head protected by prosternal lobe 7

7. Second elytral stria terminating at about basal one third of elytra Elsianus,

Second elytral stria complete or nearly so 8

8. Prothorax with a sublateral carina extending from base to apex 9

Prothorax without a sublateral carina or with a very short one 13

9. Median longitudinal groove or impression present 10

Median longitudinal groove or impression absent 11

10. Transverse impression on apical one third; two oblique subbasal impressions,

Microcylloepus,

Transverse impression absent; no oblique subbasal impressions Cylloepus,

11. No transverse or other impressions between sublateral carinae Limnius,

Transverse impression present 12

12. Impression a little anterior of middle Neoelmis,

Impression median Heterclmis,

13. Sublateral carinae very short, from one third to one half the length of thorax,

Heterlimnius,

Sublateral carinae absent Simsonia,

Groups of Stenelmis

An examination of our species shows that they may be separated
into three fairly well-defined groups as follows:

PAQB

1. First elytral stria incomplete, terminating shortly behind scutellum; granules of head
and legs elongate Nubifera group, 661

First elytral stria complete from base to apex; granules of head and legs rounded. ... 2

2. Last tarsal segment distinctly longer than the four preceding combined, the last seg-
ment usually suddenly dilated beyond the middle; tarsal claws comparatively robust,

Humerosa-sinuata group, 684

Last tarsal segment never distinctly longer than the preceding segments combined, the

last segment not as noticeably dilated; tarsal claws comparatively slender,

Crenata group, 663

Nubifera Group
Stenelmis nubifera Fall

(PI. LXXX, fig. 1; PI. LXXXI, figs. 7, 19)
1901. Stenelmis nubifera Fall, Occ. Papers Cal. Ac. Sci. VIII, p. 238.

Size. Length, 2.15-2.7 mm.; width, .87-1 mm.

Form and color. Body elongate, nearly parallel, and only slightly
wider behind middle, moderately subdepressed. Color deep reddish-
brown with a broad basal testaceous area on elytra and another be-
fore the apex.

Head. Frontal band absent; head granulate with the granules
rather large, longitudinally elongate, and separated laterally by
their own transverse diameters. Antennae and palpi testaceous ; the
antennae distinctly longer than the pronotum.

Pronotum. Length, .63-. 8 mm.; width, .73-. 9 mm. Widest just
behind middle, then gradually tapering to base; slightly narrowed
before the rounded sides and sinuate before apex. Base of pronotum
distinctly wider than apex. Entire surface moderately, closely, and
evenly granulate; the granules rounded or elongate and separated

662 The University Science Bulletin

by less than their diameters. Median sulcus moderately deep, rather
suddenly narrowed to the two posterior carinae before the scutellum.
On each side of sulcus, two elongate tubercles, the posterior one of
which is distinctly carinate before and extending to the base, the
anterior one slightly prominent posteriorly and gradually reduced
anteriorly to the surface of the disc. A slight depression on each
side of the sulcus at base.

Elytra. Length, 1.6-2 mm.; width, .87-1 mm. Basal band oc-
cupying from one third to one fourth of the elytral length; the
apical one about one fourth; suture darker to the apex. Elytral
punctures very broad and deep basally, becoming reduced in size
and depth toward the apex, on the apical band of which the punc-
tures are nearly obsolete. First elytral stria incomplete, irregularly
punctured, and extending to about one fifth of elytral length. In-
tervals nearly flat except the fourth which is slightly elevated and
broadened at base.

Venter. Brown to dull grey. Granules of abdomen a little more
elongate than on pronotum, becoming smaller and fewer towards
apex. Apex of last abdominal segment with a very broad, shallow
emargination which is usually equal to twice the greatest width of
the last tarsal segment.

Legs. Hind tibiae, .65-. 8 mm. ; middle tibiae, .5-.65 mm. ; anterior
tibiae, .5-. 65 mm. The last tarsal segment perceptibly shorter than
the four preceding combined. All femora with granules rather close
and longitudinally elongate. Middle tibia of male with a row of
about ten evenly spaced small spines on the inside. Posterior tibiae
with a similar set of spines nearly equally spaced and about fifteen
in number. Femora, tibiae, and tarsi uniformly colored as the
venter.

Notes on types. This species was described from a small series
of specimens taken at Pasadena, Cal, October 31, 1892. I have
examined the type and two paratypes.

Remarks and comparative notes. In the series of specimens ex-
amined, some variation has been noted in pronotal sculpturing; the
width of the elytral bands is slightly variable, and there is some
deviation in the ratio of the elytral length to the combined width.
The demarkation between the bands is usually indistinct. In some
specimens there is a tendency for the antennae to become darker
tow^ard the distal segment.

Notes on distribution. In addition to the types from Pasadena,
Cal., the species has been examined from the following localities:

Sanderson: Species of Stenelmis 663

■ California: Siskiyou Nat. Forest, 7-14-35, R. H. Beamer.

Oregon: Bonneville, 7-3-35, R. H. Beamer.

AVashington: Kalama, 7-4-35, R. H. Beamer.

Location of types. Type and paratypes in the collection of Mr.
H. C. Fall, Tyngsboro, Mass.; one male paratype in the Francis
Huntington Snow entomological collection. University of Kansas.

The distribution of this species has been extended for nearly
900 miles from the type locality.

Crenata Group

Key to the Species of the Crenata Group page

1. Each elytron with three longitudinal vittae sexlineata n. sp., 663

Each elytron with not more than one vitta or elytron bimaculate 2

2. Humeral spot or vitta on inside of sixth interval 3

Humeral spot or vitta embracing umbone of elytra 5

3. Body very robust, and with the elytral spots or stripe wider, covering considerably
more than the fifth interval; third elytral interval sharply elevated at base,

crenata (Say), 665
Body very elongate, with the elytral spots or stripe narrower, covering but little more
than the fifth interval; third interval but slightly elevated at base and very short, 4

4. Length 2.85-2.9 mm.; median lobe of aedeagus* distinctly constricted at middle,

exigua n. sp., 669
Length 3.2-3.4 mm.; median lobe of aedeagus more nearly parallel ... beamen n. sp., 571

5. Vitta very broad and covering nearly all of the inner space between the first and
sixth intervals ; vitta extending cephalad outside elevated apex of sixth interval,

lateralis n. sp., 672
Vitta narrower and never extending internally beyond the second or third intervals. .6

6. Lower margin of last tarsal segment with a conspicuous angular process ; species
generally larger, 3.2-3.6 mm 7

Lower margin of last tarsal segment without .svich a process; size usually smaller,
2.6-3.2.5 mm 8

7. Apical abdominal emargination equal to width of last tarsal segment; tibiae

testaceous only at base; pronotum generally rounded at sides concinna n. sp., 674

Apical emargination very inconspicuous and much less than width of last tarsal seg-
ment ; tibiae and apices of femora testaceous tarsalis n. sp., 675

8. Basal tubercle of pronotum elongate and carinate 10

Basal tubercle just perceptibly elongate and never carinate 9

9. Each elytron distinctly bimaculate knobeli n. sp., 677

Each elytron with an entire vitta hicarinata Lee, 679

10. Elytra twice longer than its width, the legs entirely testaceous exilis n. sp., 680

Elytra less than twice its width, the legs wholly or in some part dark.. /nero n. sp., 682

Crenata Group

Stenelmis sexlineata n. sp.

(Pl. LXXX, fig. 2; PI. LXXXI, fig. 23)

Size. Length, 3.2-3.6 mm.; width, 1.25-1.4 mm.
Form and color. Body elongate, parallel, moderately subde-
pressed. Color of elytra dark brown to black, and with three

* In this paper the term aedeagus is used in its less restricted sense.

664 The University Science Bulletin

longitudinal brownish testaceous vittae on each elytron. The vitta
of the fourth and fifth intervals not extending laterad of the sixth
interval to embrace the umbone.

Head. Granulations between eye and band rounded and gener-
ally separated from two to four times their own diameters. An-
tennae and palpi brownish-testaceous. Antennae distinctly shorter
than pronotal length.

Pronotum. Length, 1-1.15 mm. ; width, .95-1.12 mm. Disk of pro-
notum deep brown, but with posterior four fifths of sulcus and sur-
face caudad of lateral oblique impression, grey. Widest behind
middle, then tapering to base, but not suddenly. Rather sharply
sinuate before rounded sides, then straight and nearly parallel to
apex. Base of pronotum distinctly and conspicuously wider than
apex. Granules between the median sulcus and the lateral tu-
bercles ; fine granules separated by nearly twice their own diameters.
Median sulcus deep and extending from apical one fifth to base,
sides parallel. An oblicjue impression on cither side of median sul-
cus deep and clearly separating lateral tubercles; basal tubercle
prominent, elongate, rounded anteriorly, distinctly narrowed and
carinate posteriorly to base of pronotum. Anterior tubercle promi-
nent, rounded, scarcely wider in any one diameter. Anterior angles
of pronotum dull rufous.

Elytra. Length, 2.3-2.75 mm.; width, 1.25-1.4 mm. Each elytron
with three longitudinal browaiish-testaceous vittae. Vitta on inside
of the sixth interval entire, not at all interrupted at middle, and
occupying all of fourth and fifth intervals from base of elytron to
apex; this vitta wider at base, narrowed toward middle, and again
wider near the apex in conformity with combined widths of tiie in-
tervals covered. First interval dark brown from behind scutellum
to near apex; the second, including striae on either side, testaceous
from near its base for three fourths the length of interval. A third
vitta extending from the humerus on the outside of the sixth inter-
val to the apex. First elytral stria continuous from elytral base
to apex. Third interval distinctly raised at base and very convex.

Venter. General color grey, with the lateral and posterior margins
of abdominal segments and median portion of metasternum brown-
ish. Emargination of last abdominal segment just less than greatest
width of last tarsal segment.

Legs. Hind tibiae, .9-1.12 mm.; middle tibiae, .75-. 9 mm.; an-
terior tibiae, .85-. 95 mm. Last tarsal segment distinctly shorter
than the four preceding combined. Granules of hind femora sepa-

Sanderson: Species of Stenelmis 665

rated from two to four times their own diameters. All femora darker
above than below, the apices of femora, tibiae, and tarsi brownish-
testaceous.

Notes on types. Holotype male, 7-19-33; allotype female, 7-22-
32; and numerous paratypes, Lawrence, Kan., collected at light
from July 11 to August 5, 1930-1935, Milton W. Sanderson. Addi-
tional paratypes, Douglas county, Kansas, 9-24-1923, R. H. Beamer;
August, E. S. Tucker; Olathe, Kan., 5-6-34, S. Clare; Manhattan,
Kan., April 16, 1933, and 1934, C. W. Sabrosky; 3 miles north
Onaga, Kan., 8-15-01, French creek; Belvidere, Kan., 7-2-04; Riley
county, Kansas, September 29, C. D. Adams; Kan. (Coll. Hubbard
and Schwarz) ; Texas, Belfrage; Canyon, Tex., 7-9-33, W. Benedict.
Types and paratypes deposited in the Francis Huntington Snow
entomological collection. University of Kansas. Paratypes in the
Harvard College collection; in the collections of Dr. Paul N. Mus-
grave, Fairmont, W. Va.; and Mr. C. W-. Sabrosky, East Lansing,
Mich.

Remarks and comparative notes. The present species is the only
known one in this group which is six vittate, and is to be associated
with S. crenata (Say). It is easily separated from this, and other
related species, by the type of its maculation. The median lobe of
the aedeagus is without lateral processes as in S. crenata. An oc-
casional specimen shows very clearly the grey area between the
basal pronotal tubercles as in some specimens of S. crenata and S.
beameri n. sp.

Stenelmis crenata (Say)

(PI. LXXX, fig. 3; PI. LXXXr, fig. 20)

1824. E. creriatus Say. App. Vol. II, Keating's Exp. to source of St. Peters Riv. under
Maj. Long, Phila., p. 275.

1859. Stenelmis sordida Motschulsky. Etudes Entomologiques 8 me annee 1859 Helsingfors,
p. 51.

1869. E. crenatus Say. A description of the insects of North America, by Thomas Say,
edited by John L. LeConte. Vol. I, p. 181 (contains original description).

ORIGINAL DESCRIPTION

"Thorax with four elevated lines; each elytrum with two dull
rufous spots.

"Inhabits Pennsylvania.

"Parnn-s crenatus? Kjioch in Melsh. Catal.

"Body blackish-brown; front with two dilated, cinereous, longi-
tudinal lines; antennae and mandibles rufous; thorax with four
obtuse, elevated, longitudinal lines; two intermediate ones nearly
confluent at each end; lateral ones more distant, slightly interrupted

666 The University Science Bulletin

behind the middle; elytra with striae of dilated impressed punc-
tures; an elevated line from the humerus terminates rather before
the tip; another elevated line nearer the margin also originates at
the humerus and becomes obsolete before the middle; a third ele-
vated line originates at the middle of the base and also becomes
obsolete before the middle of the elytrum ; an oblong rufous spot on
the humerus and another near the tip; tarsi dull rufous.
"Length less than three twentieths of an inch."

writer's description

Size. Length, 3-3.35 mm.; width, 1.2-1.37 mm.

Form and color. Body generally very broad and robust, subde-
pressed, distinctly wider behind. Color of elytra dark brown to
black, each elytron with two testaceous spots or with an entire
vitta which is on the inside of the sixth interval.

Head. Granulations between eye and band rounded, separated
from one to several times their own diameters. Antennae and palpi
brownish-testaceous. Antennae distinctly shorter than pronotum.

Pronotum. Length, .87-1.1 mm.; width, .87-1.12 mm. Anterior
angles and margin rufescent. Base of pronotum from posterior
angles to anterior end of basal tubercle and all of disk between
tubercles a light grey in color. Margins and posterior half of
median sulcus generally of same color. Remainder of pronotum
dark brown to black. Pronotum widest just behind middle, then
distinctly rounded and narrowed to posterior angles; in some speci-
mens, slightly sinuate before hind angles. Base of pronotum nar-
rower than behind middle. Margins in front of rounded sides
sinuate, then faintly inwardly curved to apex. Base of pronotum
distinctly conspicuously wider than at apex. Granules between
lateral tubercles and sulcus very fine and sparse, separated by sev-
eral times their diameters. Median sulcus very deep, widest before
middle, then narrowed behind. Sulcus extending posteriorly from
apical one fourth or one fifth to base. Oblique impression on either
side of median sulcus distinct and clearly separating lateral tuber-
cles. Basal tubercle elongate, narrowed posteriorly, and distinctly
carinate to base of pronotum; anterior tubercle prominent, a little
longer than wide.

Elytra. Length, 2.12-2.37 mm.; width, 1.2-1.37 mm. Each ely-
tron with a humeral and subapical spot or with spots very distinctly
united into a vitta, but which is always on the inside of sixth in-
terval. Vitta or spots usually covering fourth and fifth intervals
with apex of vitta or subapical spot usually reaching to about one

Sanderson: Species of Stenelmis G67

half the distance between end of carina of sixth interval and apex
of elytron. First elytral stria complete from base to apex. Third
interval distinctly raised and rounded at base, the elevation ex-
tending a little beyond basal declivity. Elytral punctures deep on
disk, then smaller toward apex, but never obsolete in apical region.

Venter. Grey to brownish with apical margins of abdominal
segments usually lighter. Abdominal emargination very incon-
spicuous and considerably less than the greatest width of last tarsal
segment.

Legs. Hind tibiae, .87-1 mm.; middle tibiae, .67-. 85 mm.; an-
terior tibiae, .75-. 9 mm. Last tarsal segment equal to the combined
length of the four preceding. Granules of femora rounded, separated
from once to twice their own diameters. Femora dark except apex,
which is testaceous. Tibiae and tarsi testaceous, the tarsi slightly
darker.

Notes on types. Since the original types of this species are lost,
new types are hereby designated.

Neoholotype, neoallotype, and neoparatypes, Guthrie, Tenn., 7-
26-34, M. W. Sanderson, collected in Red river. Additional neo-
paratypes from the following localities: Hull, Que., 27, VHI, 1931,
W. J. Brown (Can. Nat. Coll.) ; Knowlton, Que., 12, VI, 8, VHI,
21, Vni, 1930, L. J. Milne (Can. Nat. Col.) ; Wakefield, Que., 7-
VIII, 31 (Can. Nat. Coll.) ; Winchester, Va., 7-21-34, M. W. San-
derson, Hogue Creek; Buffalo, N. Y.; "Mass."; Romney, N. H., VII,
18, '30, Darlington; Plymouth, VIII, 20, 1925, Darlington; Freeville,
N. Y., 6 V, 1915 (H. H. Knight Coll.) ; "Pa." (H. H. Knight Coll.) ;
Biol. Field Sta., Ithaca, N. Y., X, 2-13 (H. H. Knight Coll.) ; The
Cove, Ithaca, N. Y., X, 2, 1913 (H. H. Knight Coll.) ; Inlet Valley,
Ithaca, N. Y., X-2, 1913 (H. H. Knight Coll.) ; Ithaca, N. Y. (H. H.
Knight Coll.) ; Concordville, Md., 7-18-34, M. W. Sanderson; Alpena,
Mich., Oct. 5, '08, Nason (C. A. Frost Coll.) ; the following neopara-
types from the collection of P. N. Musgrave: Greenbrier, Tenn., June
12, '31, Little Pigeon river; Fairmont, W. Va., Aug. 3, at light; Btn.
Va.; Head and Marlinton, W. Va., July 13, Elk river; Mineral
county, W. Va., July 20, Johnny Cake Run; Mt. Storm, W. Va.,
Aug. 19, 1930, Johnny Cake Run; Wardensville, W. Va., July 19
and August 23, at light, and in Moore's Run; Huntsville, Ala., 6-17,
Dry Fork; Pocahontas county, W. Va., June 29, Minnehaha Springs.

Remarks and comparative notes. In his original description, Say
states, "Inhabits Penn." LeConte (1852) subsequently mentions
Niagara river and again Pennsylvania as localities from which speci-

668 The University Science Bulletin

mens were taken. Horn (1870), in regard to this species, states,
"Our common eastern species easily known by its broader thorax
with more strongly rounded sides. Disk is channeled and on each
side three obtuse tubercles. Entire surface is opaque and its general
aspect more depressed. Abundant in waters of Pennsylvania."

Since Say's types of »S. crcnata are apparently lost, the author has
redescribed what he regards as this species, and has designated new
types. An examination of specimens determined by LeConte and
Horn as S. crenata, examination of material from the Niagara and
Pennsylvania, and many additional specimens from a wide range of
localities over the eastern United States, shows that we have one
very common species which might easily have been that of Say.

An examination of the type female of *S. sordida Mots, shows that
it is the same species as S. crenata. Since the latter has priority by
thirty-five years, S. sordida is to be dropped from our lists. The
elytral spots are not evident in the type due to a thin incrustation of
debris. The sordid aspect of the specimen no doubt suggested its
name to Motschulsky. As indicated in the original description, his
specimen was from Pennsylvania.

Hundreds of individuals of the present species have been examined
from localities, as already indicated, and there is considerable varia-
tion within the series. There is some variation in size, relative pro-
portions of length and width of pronotum, although the two measure-
ments are generally the same, and in the character of the elytral
maculae. Without exception, the macula is confined to the inside
of the sixth interval, although the variation from an entire to a
broken vitta is notable. In some specimens there is no indication
of a separation of the humeral from the subapical spot.

Of the many specimens that have been examined, a single example
has been found in which the lower margin of the last tarsal segment
is prolonged nearly as in S. tarsalis n. sp. and S. concinna n. sp. The
latter species, however, have the humeral spot covering the umbone.
The present species maj' be separated from its closest ally, S. beameri
n. sp., by its less elongate and more depressed form, the wider vittae,
which in S. beameri usually cover no more than the fifth interval,
and by its peculiar type of genitalia. The median lobe possesses
two rounded lateral processes on the apical half, a character which
is found in no other member of the Crenata group.

Notes on distribution. In addition to the specimens from various
places designated as types, specimens from the following localities
have been examined:

Sanderson: Species of Stenelmis 669

United States:

Connecticut: Cornwall.

Massachusetts: Tyngsboio; Wellcsley; Walpole; Plymouth.

Maine: Monmouth; Paris.

New York : Batavia ; Staten Island.

Tennessee: "Tenn."

Maryland: "Md."

Pennsylvania: "Penn."; "Pen."; Lancaster.

Texas : Columbus.

District of Columbia : Washington.

Virginia: Fredrkbg.; Pennington Gap.

Michigan: Gd. Ledge; Cheboygan Co.

Kansas : Manhattan ; Lawrence ; Olathe.

Can.ad.a :

Quebec: Hull; Ivnowlton; Knowlton's Landing; Wakefield; Otter Lake;

Fairy Lk. ; South Bottom ; MacDonald ; Covey Hill ; Kazubazua.
Ontario: Bothwell; Normandale; Miner's Bay; Dundas; Walsh; Ron-

teau Pk.; Rideau R. ; Ottawa.
New Brunswick : Boiestown.

Location of types. Neoliolotype, neoallotype and neoparatypes
deposited in the Francis Huntington Snow entomological collection,
University of Kansas. Additional neoparatypes deposited in nearly
all the collections given under acknowledgments.

Stenelmis exigua n. sp.

(PI. LXXX, fig. 4)

Size. Length, 2.85-2.9 mm.; width, 1-1.1 mm.

Form and color. Body elongate, nearly parallel, scarcely wider
behind, moderately subdepressed. Color deep brown to black. Each
elytron with a narrow testaceous humeral spot and a subapical one.
Humeral spot on inside of sixth interval.

Head. Granulations between frontal band and eye minute,
rounded, and generally separated from two to four times their own
diameters. Antennae and palpi testaceous. Antennae just per-
ceptibly longer than pronotum.

Pronotum. Length, .87-. 9 mm.; width, .75-. 85 mm. Wider behind
the middle, then a little narrowed to base. But faintly sinuate be-
fore hind angles. Margin convergent from rounded sides to apex,
before which there is a double sinuation, the one immediately before
the anterior angles sm'aller and less distinct. A little wider at base
than at apex. Granules on either side of median sulcus at base fine
and separated by two or three times their diameters. Median sulcus
deep and extending from apical one fourth to base, the sides of
sulcus slightly narrowed behind. Oblique impression on either side
of median sulcus rather deeply separating lateral tubercles. Basal

670 The University Science Bulletin

tubercle rounded anteriorly and elongate, but scarcely distinctly
carinate. Anterior tubercle rounded, scarcely prominent. Anterior
angles and margin faintly to distinctly rufescent; remainder of
pronotum dull black and in some individuals with a dull greenish
cast.

Elytra. Length, 2.05-2.15 mm. Humeral spot narrow, occupying
fifth interval, extending from behind humerus to about one fifth of
elytral length. Subapical spot extending but little beyond apex of
carina of sixth interval, covering all of fifth and a part of fourth
intervals. First elytral stria complete from base to apex. Third
interval faintly elevated at base. Strial punctures moderate, never
obsolete near apex.

Venter. Dark through a light brown to dull grey or green with
posterior and lateral margins of segments lighter. Apical emargina-
tion nearly equal to width of last tarsal segment.

Legs. Hind tibiae, .87-.9 mm.; middle tibiae, .7-.8 mm.; anterior
tibiae, .75-.85 mm. Last tarsal segment a little shorter than the four
preceding combined. Granules of femora (outer side) separated by
no more than their own diameters, frequently less than. Femora
uniformly dark except apex, which is testaceous. Tibiae and tarsi
testaceous.

Notes on types. Holotype male, and allotype female, Polk, Ark.,
July 21, 1928, L. D. Beamer, collected at light. Paratype male,
same data; paratype, female, "Mo.," Otto Lugger collection.

Remarks and comparative notes. This species is very closely re-
lated to S. hea^neri n. sp., and may be separated as follows: By
actual measurement the pronotum and elytra are shorter in S.
exigua, and the legs comparatively shorter. In the few specimens of
the present species at hand, there is a fairly distinct double sinua-
tion of side of pronotum in front of rounded side, while S. beameri
has only the sinuation immediately in front of rounded sides. The
sinuation before the posterior angle of pronotum is also more distinct
in S. exigua. Reference to the male genitalia shows a much greater
constriction of the median lobe in S. exigua than in ^S. beameri. This
species is about the size of S. exiles n. sp., and S. knobeli n. sp., but
differs from either by having the humeral spot of elytra on the in-
side of sixth interval.

Location of types. Holotype and allotype in the Francis Hunt-
ington Snow entomological collection. University of Kansas. One
paratype in collection of Dr. Paul N. Musgrave. The second para-
type deposited in the collection of the University of Minnesota,
from which it was borrowed for study.

Sanderson: Species of Stenelmis 671

Stenelmis beameri n. sp.

(PI. LXXX, fig. 5)

Size. Length, 3.2-3.4 mm.; width, 1.2-1.25 mm.

Form and color. Body elongate, nearly parallel, scarcely wider
at all behind, moderately subdepressed. Color, black, each elytron
with a narrow testaceous vitta, broadly interrupted at middle. Vitta
not extending beyond the sixth interval.

Head. Granulations between frontal band and eye rounded and
generally separated by a little more than their diameters. Antennae
and palpi testaceous; antennae as long as pronotum.

Pronotum. Length, .95-1.05 mm.; width, .9-1 mm. Color, brown
to dull black. Distinctly wider behind middle, then gradually taper-
ing to base. Sinuate before the rounded sides, then convergently
straight to apex. Base of pronotum distinctly wider than apex.
Granules on either side of sulcus at base separated from slightly less
to a little more than their own diameters. Median sulcus deep and
extending from apical one fourth to base; sides parallel. An oblique
impression on either side of the median sulcus clearly separating the
lateral tubercles; the basal tubercle prominent, elongate, narrowed
toward base of pronotum, and distinctly carinate; the anterior one
prominent, rounded though very slightly longer in its longitudinal
diameter. Anterior angles and margin and posterior angles, dull
rufous.

Elytra. Length, 2.3-2.5 mm.; width, 1.2-1.25 mm. Humeral spot
narrow, occupying no more than fifth interval, and extending from
just behind the humerus to about one fourth or one third of re-
mainder of fifth interval. First elytral stria complete. Elytral
punctures moderately impressed, a little larger on disk, punctures
becoming smaller toward apex, but never obsolete before reaching it.
Third interval but slightly raised and rounded on basal declivity.

Venter. General color grey with lateral margins of metasternum
and lateral and apical margins of abdominal segments testaceous.
Emargination of last abdominal segment equal to the greatest width
of last tarsal segment.

Legs. Posterior tibiae, 1-1.1 mm.; middle tibiae, .8-. 87 mm.; an-
terior tibiae, .87-1 mm. Last tarsal segment shorter than the four
preceding combined. Granules of femora rounded and separated
by no more than their own diameters. Femora darker above and
below; the apices and margins, tibiae; and tarsi brownish-testaceous.

Notes on types. Holotype male, allotype female, and numerous
paratypes, Berryville, Ark., 7-4-34, R. H. Beamer, and M. E. Grif-

672 The University Science Bulletin

fith, collected at light. Additional material from Noel, Mo., 7-19-24,
Beamer and Lawson, at light ; Hollister, Mo., 7-28-84, M. W. Sander-
son, Long Creek.

Remarks and comparative notes. In S. beameri n. sp. there is
some variation in the length of the elytral spots, there being a tend-
ency for union in the middle of elytron in some specimens, though
there is always some clouding medially. The female allotype shows
a greyish band at base of pronotum which extends from hind angles
to anterior end of basal tubercles, then gradually curves to the op-
posite and basal angle of pronotum. This also occurs in some other
specimens of this species. This coloration is quite characteristic of
S. crenata (Say).

The present species keys out with S. crenata by its type of macula-
tion, but is readily separated by its much more slender and elongate
form. The maculation of S. beameri scarcely covers more than the
fifth interval, the raised third interval at base is slight, compared
with S. crenata, and the median lobe of the aedeagus of S. beameri is
without lateral processes.

Location of types. Holotype, allotype, and paratypes deposited
in the Francis Huntington Snow entomological collection, Univer-
sity of Kansas. Additional paratypes in the collections of Dr. Paul
N. Musgrave, Fairmont, W. Va. ; and Harvard College.

Stenelmis lateralis n. sp.

(PI. LXXX, fig. 6; PI. LXXXI, figs. I, 15, 21)

Size. Length, 2.65-3 mm.; width, .95-1.1 mm.

Form and color. Body elongate, sides parallel, scarcely, if at all,
wider behind than across humeri; moderately subdepressed. Color,
black ; each elytron with a very broad, yellow-testaceous vitta, which
embraces umbone and extends on inside of sixth interval nearly to
apex. Vitta also extending cephalad on lateral side of sixth interval.

Head. Granulations between eye and band, separated by about
twice their diameters. Antennae and palpi yellow testaceous; the
antennae slightly longer than pronotum.

Pronotum. Length, .83-. 87 mm.; width, .75-. 8 mm. Widest be-
hind middle, then gradually tapering to base, before Avhich tlicre is a
slight sinuation; lateral margin sinuate in front of rounded sides,
then narrowed and straight to apex. Base slightly wider than apex.
Granulations on either side of median sulcus at base separated by
two to three times their own diameters. Sulcus moderately deep and
extending from apical one third to near the base, perceptibly nar-
rowed behind. Oblicjue impression on either side of sulcus, distinct

Sanderson: Species of Stenelmis 673

and clearly separating lateral tubercles. Basal tubercle elongate,
narrowed toward base of pronotum, but not distinctly carinate; the
anterior one rounded ; entire anterior margin of pronotum rufescent,

Elytra. Length, 1.95-2.15 mm.; width, .95-1.1 mm. Vitta oc-
cupying all of inner space between first stria and sixth interval.
Basal margin of elytra darker. Sutural stripe often a little broader
toward apex. First elytral stria complete from base to apex. Third
interval perceptibly raised and rounded basally. Elytral punctures
large and deep, becoming smaller and shallower tow^ard apex;
punctures obsolete behind end of carina of sixth interval.

Venter. General color brownish to grey, the abdominal segments
with posterior border testaceous. Apical emargination about equal
to width of last tarsal segment.

Legs. Hind tibiae, .8-. 85 mm.; middle tibiae, .65-. 7 mm.; anterior
tibiae, .65-.7 mm. Last tarsal segment distinctly shorter than four
preceding combined. Granules of femora closely placed and usually
separated by less than their own diameters. All femora darker above
and below, the tibiae and tarsi testaceous.

Azotes on types. Holotype male, allotype female, and numerous
paratypes from Berryville, Ark., 7-4-34, R. H. Beamer and M. E.
Griffith, collected at light. One paratype from Schellburg, Pa.,
VIII 16. Additional material from Hollister, Mo., 7-28-34, M. W.
Sanderson, Long creek; Noel, Mo., 7-19-24, Beamer and Lawson,
at light; Ireland, Miss., 7-7-34, M. E. Griffith, at light; Lucedale,
Miss., June 22, P. N. Musgrave, White's creek; Winchester, Va.,
7-21-34, M. W. Sanderson, Hogue creek; Clarksville, Tenn., 1915,
at light.

Remarks and comparative notes. The present species is of par-
ticular interest, since it appears to be very abundant in several lo-
calities of the Ozark Mountain region in Arkansas and Missouri.
The elytral vitta is always entire and is much broader than in any
other known species of our fauna. The extension of the vitta
cephalad on the outside of the carina of the sixth interval is very
characteristic of this species. This character is found in an oc-
casional specimen of »S. mera n. sp., but the broader stripe with no
indication of bimaculation will readily distinguish the two.

Location of types. Holotype, allotype, and paratypes in the
Francis Huntington Snbw entomological collection, University of
Kansas. Additional paratypes in the collections of Dr. Paul N.
Musgrave, Canadian national collection. Harvard College collec-
tion, C. A. Frost collection, and the collection of Harry H. Knight,
Iowa State College.
43—4141

674 The University Science Bulletin

Stenelmis concinna n. sp.

(PI. LXXX, fig. 7)

Size. Length, 3.3-3.6 mm.; width, 1.3-1.5 mm.

Form and color. Body generally broad and robust, moderately
convex, nearly parallel behind. Color, dark brown to dull black;
each elytron with a testaceous spot on humerus which extends on
inside of sixth interval, and an elongate subapical one.

Head. Granulations between eye and band rounded and sep-
arated by about twice their own diameters. Antennae and palpi
brownish-testaceous. Antennae equal in length to pronotum.

Pronotum. Length, 1-1.1 mm.; width, .9-1 mm. Widest just be-
hind the middle, then rounded to posterior angles. In some speci-
mens slightly sinuate before hind angles. Margins in front of
rounded sides narrowed and subparallel to apex. Before the middle
of sides, a slight sinuation. Base of pronotum distinctly wider than
apex. Granules between lateral tubercles and sulcus separated by
about twice their own diameters. Median sulcus very deep, margin
behind middle conspicuously raised and prominent. Inner sides of
sulcus perpendicular in some specimens just before base; sulcus ex-
tending posteriorly from apical one fourth and slightly narrowed
toward base. Obliciue impression on either side of sulcus distinct
and rather deeply separating lateral tubercles. Basal tubercle elon-
gate, narrowed toward base and distinctly carinate to base, in some
individuals acutely so; anterior tubercle elongate, about three times
longer than its greatest width ; anterior angles and margin rufescent.

Elytra. Length, 2.35-2.65 mm.; width, 1.3-1.5 mm. Small humeral
spot embracing umbone, extending to and covering fourth interval
at base and occupying only about one fifth of elytral length. Sub-
apical spot extending from a point considerably behind the middle
to a little beyond the apex of carina of sixth interval and occupying
fifth and a part of fourth and sixth intervals. First elytral stria
complete from base to apex. Third interval distinctly elevated and
rounded at base, the elevation extending to near the posterior border
of humeral spot. Elytral punctures deep at base and on disk and
becoming shallow and smaller toward apex, never obsolete in apical
region.

Venter. General color deep brown to grey, with apical margins
of abdominal segments lighter. Last ventral abdominal segment
almost wholly dark. Apical emargination conspicuous, equal to
width of last tarsal segment.

Legs. Hind tibiae, 1-1.12 mm.; middle tibiae, .87-.95 mm.; an-

Sanderson: Species of Stenelmis 675

terior tibiae, .9-1 mm. Last tarsal segment shorter than the four
preceding. Lower margin of apex of this segment prolonged into a
conspicuous angulate process. Granules of hind femora rounded,
very closely placed. Femora and tibiae dark in color, except base
of tibia, which is brownish-testaceous. Tarsi brownish-testaceous.

Notes on types. Holotype male, allotype female and a number of
paratypes. North River, N. Y., 7-7-34, M. W. Sanderson, collected
in Hudson river. Additional paratypes, Wakefield, Quebec, May
29-Aug. 7, 1930-1932, W. J. Brown (Can. Nat. Coll.) ; Knowlton,
Quebec, June 29-Aug. 8, 1928-1930, L. J. Milne (Can. Nat. Coll.) ;
Bedford, Mass., VII-15-11 (C. A. Frost Coll.) ; mouth of Williams
river, W. Va., July 10, 1933 (P. N. Musgrave Coll.) ; Marlin, W. Va.,
July 29, 1930, Greenbrier river (P. N. Musgrave Coll.) ; "N. C."

Remarks and com-parative notes. In the large series of this species
which the author has examined, all of the specimens show a distinct
and wide separation of the humeral from the subapical spot, there
being no indication of a union. A single specimen of this species
from Marlin, W. Va., does not have the process of the tarsal seg-
ment prominent. However, the humeral spot covers the humerus,
and the median lobe of aedeagus is without processes — characters
which exclude it from being placed with &. crenata (Say).

This species is very closely related to S. tarsalis n. sp. by the pres-
ence of the angulate process on the lower margin of last tarsal seg-
ment, and by its type of elytral maculation. It may be distinguished
from S. tarsalis by having the tibiae nearly completely dark; the
apical abdominal emargination is equal to the tarsal width, and the
humeral spot is much shorter than in S. tarsalis. The sides of the
median lobe are nearly parallel in the present species.

Location of types. Holotype, allotype, and paratypes in the
Francis Huntington Snow entomological collection, University of
Kansas. Additional paratypes in the Canadian national collection,
C. A. Frost collection and the collection of Dr. Paul N. Musgrave.

Stenelmis tarsalis n. sp.

(PL LXXX, fig. 8; PI. LXXXI, figs. 13, 18)

Size. Length, 3.2-3.5 mm.; width, 1.25-1.5 mm.

Form and color. Very broad and robust, moderately convex, wider
behind than across the humeri. Elytra black; each elytron with a
small entire spot which covers humerus, and an elongate subapical
one.

Head. Granulations between eye and band rounded, and gen-
erally separated by two to three times their own diameters. An-

676 The University Science Bulletin

tennae and palpi brownish-testaceous, antennae slightly shorter than
pronotal length.

Pronotum. Length, .95-1.12 mm.; width, 1-1.2 mm. Widest con-
siderably behind the middle, then rounded to posterior angle, before
which there is a vague sinuation. Lateral margins narrower in front
of rounded sides and nearly straight to apex ; a very feeble sinuation
about half way between rounded sides and the anterior angles. Base
of pronotum distinctly wider than apex. Granules on each side of
sulcus separated by from once to twice their own diameters. Median
sulcus very deep and with lateral margins perpendicularly produced
internally for nearly the entire length. Sulcus extending posteriorly
from about the apical one fifth to near posterior margin of pronotum,
nearly parallel-sided throughout its length. A distinct oblique im-
pression on either side of median sulcus clearly separating lateral
tubercles. Basal tubercle elongate, narrowed posteriorly, and dis-
tinctly carinate almost to basal margin of pronotum, anterior one
elongate, nearly three times longer than its own width. Anterior
margin of pronotum rufescent, more distinctly so at angles. A light
greyish area between the basal tubercles.

Elytra. Length, 2.25-2.6 mm.; width, 1.25-1.5 mm. Small humeral
spot embracing umbone, extending posteriorly on inside of sixth in-
terval for about one fourth of elytral length. Spot usually not ex-
tending beyond fifth interval internally. Subapical spot extending
from behind middle to a point slightly beyond end of carina of sixth
interval. This spot embracing fifth and part of fourth intervals in
this region. First elytral stria complete from base to apex. Third
interval distinctly elevated and rounded on basal declivity. Elytral
punctures very broad and deep on either side of third interval at
base. Remaining punctures deep, but becoming shallow and smaller
toward apex of elytron, never obsolete in apical region.

Venter. General color, grey to nearly light blue, sometimes iri-
descent, and with apical margins of abdominal segments and nearly
the entire apical segment brownish-testaceous. Apical emargina-
tion of last segment inconspicuous, not more than one half the
greatest width of last tarsal segment.

Legs. Hind tibiae, .87-1.12 mm.; middle tibiae, .75-. 9 mm.; an-
terior tibiae, .82-. 95 mm. Last tarsal segment shorter than the four
preceding combined. Granules of hind femora separated by little
more than their own diameters. All femora entirely grey in color
except apices, which are lighter. Tibiae brownish-testaceous, but
with tarsi just slightly darker. Lower margin of last tarsal segment
prolonged into a prominent process.

Sanderson: Species of Stenelmis 677

Notes on types. Holotype male, allotype female, and a number
of paratypes, Winchester, Va., 7-21-34, M. W. Sanderson, Hogue
creek. Additional paratypes: Berryville, Ark., 7-4-34, R. H.
Beamer, at light; Ottawa county, Okla., June 4, 1930, 7-30-34, M. W.
Sanderson; Guthrie, Tenn., 7-26-34, M. W. Sanderson; Annadale,
Va., 7-21-34, M. W. Sanderson; Lynchburg, Va., July 12, P. N.
Musgrave, Flat creek; Burlington, W. Va., August 22, 1930, Patter-
son creek; Wardensville, W. Va., August 22, 1933, Moore's Run;
July 19, Moore's Run, and at light; Romney, W. Va., Aug. 22, 1930,
South branch Potomac; Maysville, W. Va., Aug. 19, 1933, Lunice
creek; Pocahontas county, W. Va., July 31, 1930, Minnehaha Springs;
Franklin, W. Va., Aug. 9, 1933, Thorn creek; Clifton Forge, Va.,
July 13, Dunlap creek, P. N. Musgrave, at light; Batavia, N. Y., 7
Sept., 1915, and 8 Sept., 1916, H. H. Knight; Ottawa, Ontario, 14,
VIII, 1912, G. Beaulieu; Ottawa, Canada, 15, VIII, 1912, G.
Beaulieu.

Remarks and comparative notes. In the type there is a prominent
post-coxal elevation on the first abdominal segment, but which is
less conspicuous in other specimens. This character is also found
in other species, but usually to a less marked degree. The anterior
and the posterior lobes of the prosternum are brownish-testaceous,
a variable characteristic of this species.

This species and S. cincinna n. sp. are our largest forms of
Stenelmis in which the umbone is covered by the humeral spot.
They might easily be confused with each other, both species pos-
sessing the same type of elytral maculation. In addition, they have
the lower margin of the last tarsal segment prolonged into a prom-
inent process. The present species is to be separated from S.
concinna by its less conspicuous abdominal emargination, by its
entirely testaceous tibiae, and by the shape of its aedeagus. The
sides of the median lobe are distinctly sinuate, and not straight and
nearly parallel as in S. concinna.

Location of types. Holotype, allotype, and paratypes in the
Francis Huntington Snow entomological collection, University of
Kansas. Additional paratypes in the collection of Dr. Paul N. Mus-
grave, H. H. Knight collection, and Canadian national collection.

Stenelmis knobeli n. sp.

(PI. LXXX, fig. 9)

Size. Length, 2.75-3 mm.; width, 1-1.05 mm.
Form and color. Body elongate, moderately subdepressed, but
little wider behind. Color of elytra deep brown approaching black.

678 The University Science Bulletin

Each elytron with a humeral spot which embraces the umbone, and
a subapical spot.

Head. Granules between eye and band separated by a little more
than their own diameters. Antennae and palpi testaceous, the palpi
lighter. Antennae equal to pronotal length.

Pronotum. Length, .85-.87 mm.; width .8 mm. Wider behind the
middle, then conspicuously narrowed, but scarcely sinuate before
hind angles. Convergent from rounded sides to apex, the sides dis-
tinctly emarginate at middle. A little wider at base than apex.
Granules on either side of median sulcus at base fine, rather con-
spicuous and separated by their own diameters. Median sulcus
shallow, sides nearly parallel, extending from apical one third to
apex of basal declivity. Oblicjue impression on either side of median
sulcus rather shallowly separating lateral tubercles. Basal tubercle
broadly rounded, perceptibly elongate, and not at all carinate be-
hind. Anterior tubercle rounded, not at all prominent. Anterior
angles and margin of pronotum rufescent, the remainder of prono-
tum dull black.

Elytra. Length, 2-2.25 mm. Humeral spot covering umbone and
most of fourth and fifth intervals at basal one fifth. Subapical spot
elongate, covering most of third, fourth and fifth intervals, extending
obliquely inside to the first stria which it follows nearly to elytral
apex. Subapical spot ex-tends cephalad on the outside of carnia of
sixth interval, becoming fainter near the middle of side of elytra.
First elytral stria complete from base to apex. Third interval not
at all elevated at base. Strial punctures moderately large, less so
apically, but never obsolete near apex.

Venter. Light brown to dull grey, posterior margins and last ab-
dominal segments lighter. Apical emargination nearly equal in
width to last tarsal segment.

Legs. Hind tibiae, .85-. 9 mm.; middle tibiae, .7-. 75 mm.; anterior
tibiae, .75-.8 mm. Last tarsal segment equal to the four preceding
combined. Granules of femora (outer side) prominent and separated
by much less than their own diameters. Femora nearly uniformly
dark except at apex. Tibiae and tarsi testaceous.

Notes on types. Holotype male and allotype female, Hope, Ark.,
July 28, 1932, L. Knobel.

Remarks and comparative notes. According to the key this species
is associated with S. bicarinata Lee. by the character of the basal
tubercle of pronotum. Reference to my description of S. bicarinata
will show that S. knobeli is considerably smaller, and each elytron
is bimaculate. It bears its closest resemblance, however, to S. exilis

Sanderson: Species of Stenelmis . 679

n. sp., both in size and general appearance. From S. exilis it may
easily be distinguished by the absence of basal elevation of third
elytral interval, the absence of posterior carination of basal tubercle
of pronotum, by its darker femora, and by differences in the median
sulcus of pronotum. S. exilis lacks the extension of the posterior
elytral spot on outside of sixth interval found in the present species.
The pronotum is a little narrower in proportion to its length than
in S. exilis.

Location of types. Types located in the collection of Dr. Paul N.
Musgravc, Fairmont, W. Va.

Stenelmis bicarinata Lee.

(PI. LXXX, fig. ](); PI. LXXXI, figs, fi, 17)
1852. S. bicariiiatii>! LeConte, Proc. Ac. N. S. Pliila. VI. p. -14.

Size. Length, 2.8-3.25 mm. ; width, 1.1-1.25 mm.

Form and color. Body elongate, sides of elytra nearly parallel,
convex. Color of elytra dark brown to black, each elytron with an
entire vitta covering humerus and extending inside of sixth interval
to near the apex.

Head. Granulations between eye and band rounded, generally
separated from two to four times own diameters. Antennae and
palpi light brownish-testaceous; antennae as long as or slightly
longer than pronotum.

Pronotum. Length, .85-1 mm.; width, .8-. 9 mm. Disk of prono-
tum grey and usually with blue or greenish reflections. Widest be-
hind the middle, then convergent to base, only a faint indication of
a sinuation before the angle. Usually slightly sinuate before rounded
sides, then convergent and straight to apex. Base of pronotum wider
than apex. Granules between basal tubercle and sulcus separated
from one to three times their own diameters. Median sulcus mod-
erately deep, inner sides only occasionally perpendicular and usually
gradually sloping from margins of sulcus. Sulcus extending pos-
teriorly from about apical one fourth to near the base, somewhat
convergent on basal pronotal declivity. A moderately distinct ob-
licjue impression on either side of median sulcus shallowly separating
lateral tubercles. Basal tubercle rounded, perceptibly elongate, and
never carinate; a rather deep impression on the inside of basal
tubercle about one third from base of pronotum. Anterior tubercle
rounded, not at all prominent. Anterior angles and margin rufescent.

Elytra. Length, 2.15-2.45 mm.; width, 1.1-1.25 mm. Vitta em-
bracing umbone, extending on the inside of sixth interval, covering
all of the fourth and fifth, and terminating about half way between

680 . The University Science Bulletin

end of carina of sixth interval and apex of elytron. First elytral
stria complete from base to apex. Third interval very faintly to
distinctly raised and rounded on basal declivity. Strial punctures
prominent on disk, but becoming shallower and smaller toward apex,
never obsolete.

Venter. General color bluish-grey with posterior margins of ab-
dominal segments brownish-testaceous. Last abdominal segment in
great part testaceous, with the emargination barely less than greatest
width of last tarsal segment.

Legs. Hind tibiae, .88-1 mm.; middle tibiae, .75-. 85 mm.; an-
terior tibiae, .8-. 9 mm. Last tarsal segment shorter than the four
preceding combined. Granules of femora usually separated by less
than their diameters. Femora and tibiae grey except bases and
apices, which are brownish-testaceous. Tarsi testaceous.

Notes on type. The type of this species from Ohio has been ex-
amined.

Remarks and comparative notes. This species is closely related to
S. mera n. sp., from which it may be easily separated by its greater
length, and by having no posterior elevation of the basal tubercles
of pronotum. In addition there is no tendency toward quadri-
maculation which is often found complete in *S. mera. The median
lobe of the aedeagus is more acute and parallel in S. bicarinata Lee.
than in S. mera.

Notes on distribution.

Kansas: Lawrence, June 4-Aiig. 7, 1930-1935, M. W. Sanderson and L. S.
Henderson; Saline county, July 7, 1925, R. H. Beamer; Woodson county,
July 31, 1923, Beamer and Lawson; Riley county, Marlatt and J. B. Norton
(Kansas State College collection).

Te-xas: "Tex." Belfrage (Harvard College collection); several additional
specimens labeled "Tex." (Harvard College collection); "Tex.," May (W.
Knaus collection) ; "Tex." (Univ. of Minn, collection) ; Camp San Saba,
5-28-04 (W. Knaus collection).

New Jersey: "N. J." (W. Knaus collection); "N. J." (C. A. Frost collec-
tion).

New York: Ithaca, July 10, 1901 (H. H. Knight collection).

Vermont: Bennington county, July, 1894 (H. H. Knight collection).

Pennsylvania: "Penn." (Harvard College collection.)

Location of types. Type in the LeConte collection. Harvard
College.

Stenelmis exilis n. sp.

Size. Length, 2.8-3 mm.; width, 1-1.1 mm.

Form and color. Body elongate, moderately subdepressed, a little

Sanderson: Species of Stenelmis 681

wider behind. Color of elytra deep brown. Each elytron with a
humeral spot which embraces the mnbone, and a subapical spot.

Head. Granules between eye and band separated by their own
diameters. Antennae and palpi yellow-testaceous. Antennae equal
to the pronotal length.

Pronotum. Length, .85-. 87 mm.; width, .82-. 87 mm. Wider be-
hind middle, then conspicuously narrowed and slightly sinuate be-
fore hind angles. Convergent and nearly straight from rounded
sides of apex, distinctly sinuate at middle. Wider at base than apex.
Granules on either side of median sulcus at base very fine and in-
conspicuous. Median sulcus deep, the sides nearly parallel, extend-
ing from apical one fourth to base. A rather deep oblique impres-
sion on either side of median sulcus, clearly separating lateral
tubercles. Basal tubercle rounded, prominent, elongate, and dis-
tinctly carinate behind; anterior tubercle rounded, prominent. An-
terior angles and margin of pronotum rufescent.

Elytra. Length, 2.05-2.12 mm.; width, 1-1.1 mm. Humeral spot
covering umbone and most of fourth and fifth intervals at basal one
fifth; subapical spot elongate and terminating a little beyond the
carina of sixth interval, and also covering most of fourth and fifth
intervals. First elytral stria complete from base to apex. Third
interval elevated only on basal declivity. Elytral punctures large
and deep, never obsolete apically.

Venter. Light brown to dull grey and usually with posterior half
of abdominal segments lighter. Apical emargination equal in width
to last tarsal segment.

Legs. Hind tibiae, .IT-.Sl mm.; middle tibiae, .62-. 7 mm.; anterior
tibiae, .65-.77 mm. Last tarsal segment equal to four preceding
combined. Granules of femora separated by no more than their own
diameters. Femora and tibiae uniformly testaceous, the tarsi but
little darker.

Notes on types. Holotype female and two paratype females,
Berryville, Ark., 7-4-34, R. H. Beamer, collected at light.

Remarks and comparative notes. This species shows a striking
resemblance to some of the smaller specimens of S. beameri n. sp.,
but is at once separated by having the humeral spot covering the
umbone, and by the legs jvhich are entirely testaceous. In S. beameri
the apical one fourth only of the femur is testaceous, the tibia of
the same color.

Location of types. Types deposited in the Francis Huntington
Snow entomological collection. University of Kansas.

682 The University Science Bulletin

Stenelmis mera n. sp.

(PI. LXXX. fig. 11; PI. LXXXI, fig. 22)

Size. Length, 2.6-2.85 mm.; width, 1-1.2 mm.

Form and color. Body elongate, moderately convex. Sides of
elytra parallel and scarcely wider behind. Color of elytra dark
brown to black; each elytron with an entire vitta which embraces
the umbone and extends on the inside of sixth interval nearly to
apex; or vittae interrupted at middle to appear or nearly to appear
quadrimaculate.

Head. Granulations between eye and band rounded, generally
separated from one to three times their own diameters. Antennae
and palpi light brownish-testaceous; antennae longer than pronotum.

Pronotum. Length, .8-. 85 mm.; width, .8-. 9 mm. Disk of pro-
notum dark brown to grey and usually with a darker fuscous area
between basal tubercles; this frequently extending cephalad on
either side of median sulcus. Widest behind middle, then convergent
and straight to base. Slightly sinuate before rounded sides, then
convergent and nearly straight to apex. Granules on inside of basal
tubercles separated from once to twice their diameters. Median
longitudinal sulcus deep; the inner sides, especially near base, per-
pendicular. Sulcus extending posteriorly from apical one fourth to
base, somew^hat narrowed before basal declivity. A distinct oblicjue
impression on either side of median sulcus distinctly separating
lateral tubercles. Basal tubercle rounded anteriorly, narrowed and
elongate toward base of pronotum and distinctly carinate; anterior
tubercle rounded, scarcely prominent. Anterior angles and margin
rufescent.

Elytra. Length, 1.9-2.1 mm.; width, 1-1.2 mm. Vitta embracing
umbone and extending on inside of sixth interval, usually covering
all of fourth and fifth intervals and terminating shortly before apex.
Vitta always with a more or less distinct band or clouding medially.
First elytral stria complete from base to apex. Third interval
usually very distinctly raised at base. Elytral punctures large and
prominent and becoming smaller toward apex, never obsolete.

Venter. General color grey or brownish, with posterior margins
of abdominal segments light brownish-testaceous. Emargination of
last abdominal segment slightly more than greatest width of last
tarsal segment.

Legs. Hind tibiae, .75-. 86 mm.; middle tibiae, .62-.75 mm.;
anterior tibiae, .65-. 8 mm. Last tarsal segment shorter than the
four preceding combined. Granules on lower side of femora rounded,

Sanderson: Species of Stenelmis 683

separated by less than their diameters. Femora, except apex, gray;
tibiae variable in coloration though usually brownish-testaceous.

Notes on types. Holotype male, allotype female, and eight para-
types, Guthrie, Tenn., 7-26-34, M. W. Sanderson, collected in Red
river. Additional paratypes from East Homer, N. Y., 7-5-34, M. W.
Sanderson, East river; Ithaca, N. Y., July 10, '01; North river, N. Y.,
7-7-34, M. W. Sanderson, Hudson river; Eureka Springs, Ark.,
7-4-34, R. H. Beamer; Schellburg, Pa., VIII, 16; "Penn.," (Otto
Lugger Coll.) ; "N. C," (H. H. Knight Coll.), (Otto Lugger Coll.) ;
N. C, Mass. ; Greenbrier, Tenn., June 12, '31, P. N. Musgrave, Little
Pigeon river; South Bottom, Quebec, 16.6. 1928, G. H. Fisk (Can.
Nat. Coll.); Knowlton, Quebec, 19.6. 1928, G. H. Fisk (Can. Nat.
Coll.) ; Keyser, W. Va., 7-22-34, M. W. Sanderson, New creek.

From West Virginia, material has been collected by P. N. Mus-
grave from the following localities: Wardensville, July 19, at light;
Great Cacpon, July 24, 1933, Cacpon river; Meadow Branch of
Sleepy creek, July 23, 1933; Richmond, July 11, 1933; North Fork
Cherry; mouth of Williams river, July 10, 1933; Franklin, August 9,
1933, Thorn creek; Maysville, August 19, 1933, Lunice creek; Rom-
ney, August 22, 1930, South Branch Potomac; Burlington, August
22, 1930, Patterson creek; Stony river, August 21, 1930; Pocahontas
county, July 31, 1930, Minnehaha Springs; Big Spring creek, June
3; Fairmont, August 3; New creek, August 20, 1930.

Remarks and comparative notes. Very often specimens of S.
mera n. sp. are found in which there is a testaceous spot on the out-
side of carina of sixth interval near its apex, but which is always
short and continuous with the distal stripe. This character ap-
proaches that of &. lateralis n. sp., from which this species is easily
separated by its much narrower elytral vitta ; the vitta of &. lateralis
occupies all intervals inside of the sixth except the first. The
present species is more nearly related to &. bicarinata Lee, but is
separated by its shorter form, the elongated and carinate basal
tubercle, and the median clouding of elytral vitta which in some
instances clearly separates the humeral from the subapical spot.
In S. bicarinata Lee. the vitta is always entire with no indication
of a median cloud. >S. inera has been identified as S. linearis Zimm.
in some collections. As indicated elsewhere in this paper, this species
is a synonym of (S. humerosa Mots. It is properly placed in the
Sinuata-humerosa group, and not in the Crenata group, to which
S. mera belongs.

Location of types. Holotype, allotype, and paratypes deposited

684 The University Science Bulletin

in the Francis Huntington Snow entomological collection, Uni-
versity of Kansas. Additional paratypes in the Canadian national
collection, collections of Dr. Paul N. Musgrave; University of
Minnesota; and Dr. Harry H. Knight, Ames, Iowa.

SiNUATA-HuMEROSA GrOUP
Key to the Species of the Sinuata-Humerosa Group

PAGE

1. Femora punctulate and not at all granulate 2

Femora distinctly granulate 3

2. Longitudinal vitta of elytra complete from base to apex; processes on median lobe

of aedeagus evenly rounded douglasensis n. sp., g85

Each elytron distinctly bimaculate ; processes on median lobe of aedeagus subangulate
anteriorly grossa n. sp., 686

3. Elytra immaculate 4

Elytra maculate or vittate 6

4. Median band of head as wide as two lateral ones combined; length of body,

2.5 mm.; width, .87 mm parva n. sp., ggg

Median band but little wider than either lateral one; length of body, over 2.7 mm.;
width, over 1 mm 5

5. Length, 3.25-3.4 mm.; processes of median lobe of aedeagus present and distinct,

juscata Blatch., 689
Length, 2.7-2.8 mm. ; processes of median lobe nearly absent and very inconspicuous,

hungerfordi n. sp., 690

6. Humeral spot distinctly covering umbone 7

Humeral spot on inside of si.xth interval 8

7. Femora and tibiae entirely grey humerosa Mots., 692

Femora grey, tibiae testaceous mirabilis n. sp., 693

8. Antennae or palpi, or both, dark brown to black 9

Antennae and palpi testaceous 11

9. Last six or seven antennal segments nearly always shining black, the palpi testaceous,

antenrialis n. sp., 695
Palpi usually dark brown to piceous, the antennae nearly always lighter 10

10. Length, 2.7-3.2 mm. Lateral processes about two thirds the width of median lobe

of aedeagus quadrimaculata Horn., 696

Length, 2.4-2.65 mm., processes about one third the width of median lobe,

musgravei n. sp., 698

11. Sides of pronotum in anterior third sharply divergent sinuata Lee, 699

Sides of pronotum in anterior third parallel or convergent 12

12. Process of median lobe of aedeagus absent or very inconspicuous 13

Process of median lobe present and conspicuous 14

13. Elytral stripe present and entire; processes absent decorata n. sp., 70I

Faint humeral and subapical spots; processes very narrow and inconspicuous,

hungerfordi n. sp., 690

14. Processes of median lobe evenly rounded 15

Processes subangulate anteriorly vittipennis Zimm., 702

15. Lateral processes of median lobe as wide as lateral lobes near apex; body more
convex, the vittae less distinctly marked convexula n. sp., 704

Lateral processes about one half the width of lateral lobe near apex ; body less
convex, vittae more distinctly marked markelii Mots., 705

Sanderson: Species of Stenelmis 685

Stenelniis doiiglascnsis n. sp.

(PI. LXXX, fig. 12)

Size. Length, 3.35-3.6 mm.; width, 1.2-1.5 mm.

Form and color. Body elongate, convex, nearly parallel. Elytra
black; each elytron with a distinct longitudinal vitta on inside of
sixth interval.

Head. Granulations of head obsolete. Antennae and palpi light
brownish-testaceous. Antennae distinctly shorter than pronotum.

Pronotum. Length, 1-1.1 mm.; width, .9-1 mm. Pronotum grey
to dull black; the prominences lighter. Apical margin of pronotum
not rufescent. Sides nearly parallel in basal half and with a dis-
tinct sinuation before the hind angles; the hind angles very promi-
nent. Narrowed anteriorly, then nearly parallel or divergent in
front. Pronotum as wide at base as width behind middle; fre-
quently distinctly wider. Width at apex shorter than at base. Sur-
face of pronotum indistinctly punctulate. Median longitudinal
sulcus deeply channeled from apical one fifth to base, but little
narrowed posteriorly. Oblique impression on either side of median
sulcus veiy distinct, clearly separating lateral tubercles. Basal
tubercle is rounded anteriorly and carinate behind. Anterior tuber-
cle prominent and rounded.

Elytra. Length, 2.4-2.5 mm. Each elytron with an entire longi-
tudinal vitta extending to a point about one third of the distance
between apex of carina of sixth interval and apex of elytra; vitta
occupying nearly all of fourth and fifth intervals. Occasionally the
second and eighth intervals are faintly longitudinally vittate. First
elytral stria complete. Third interval moderately elevated at base,
the elevation extending only about one sixth of the elytral length.
Elytral punctures moderately deep on disk and not entirely obsolete
on apical declivity.

Venter. Dull green to grey or dull black. Apical abdominal
emargination a little less than width of last tarsal segment.

Legs. Hind tibiae, 1.1 mm.; middle tibiae, .87-. 9 mm.; anterior
tibiae, .9-. 95 mm. Last tarsal segment distinctly longer than pre-
ceding four combined. Granules of femora and tibiae replaced by
minute punctures. Femora and tibiae of same color as venter. The
tarsi, brownish.

Notes on types. Holotype male, allotype female, and six para-
types, Cheboygan county, Michigan, July 10, 1936, T. C. Lawrence.
Collected on submerged chunks of wood in edge of water along east
shore of Douglas Lake. Additional paratypes from Nottawa, Mich.,

686 The University Science Bulletin

Sept. 1, 1933, C. W. Sabrosky. Collected on old pieces of wood in
1 to IMi feet of water in Sand Lake. According to notes of the
collector, ''this lake has no inlet or outlet, being fed entirely by
underground springs."

Remarks and comparative notes. S. douglasensis n. sp. is closely
related only to S. grossa n. sp. by the complete absence of distinct
granules over the surface of the body. As in S. grossa, the granules
are replaced by distinct punctures. It is distinguished from S.
grossa by its entire longitudinal vittae and the evenly rounded
lateral processes of the median lobe of aedeagus. This is one of
the few species which, apparently, confines itself to the lakes of
the North. In some examples the second and eighth elytral intervals
are faintly testaceous for a part of their lengths.

Location of types. Holotype, allotype, and paratypes in the
Francis Huntington Snow entomological collection, University of
Kansas. Additional paratypes in the collections of Dr. Paul N.
Musgrave, Harvard College, and University of Michigan.

Stenelmis grossa n. sp.

(PI. LXXX, fig. 13)

Size. Length, 3.25-3.6 mm.; width, 1.25-1.4 mm.

Form and color. Body elongate, convex, nearlj^ parallel. Elytra
reddish-brown to dark grey; each elytron usually with a faint
humeral and subapical testaceous spot. Humeral spot on inside of
sixth interval.

Head. Granulation nearly absent between eye and band, but a
small group of granules near posterior end of band. Antennae and
palpi light brownish testaceous, the antennae considerable shorter
than pronotum.

Pronotum. Length. 1-1.12 mm.; width, .9-1.05 mm. Disk of pro-
notum dull grey to deep blackish-grey, the anterior part of sulcus
light fuscous. Apical margin of pronotum rufescent. Sides of pro-
notum parallel or divergent on basal half and without a sinuation
before basal angles. Narrowed anteriorly about the middle, dis-
tinctly sinuate, then sides nearly parallel or usually distinctly
divergent to apices. Pronotum as wide at the base as width behind
the middle, often distinctly wider. Width at apex shorter than at
base. Granules not evident on either side of base of median sulcus.
Median longitudinal sulcus deeply channeled from about the apical
one fourth to base, narrowed posteriorly. Oblique impression on
either side of sulcus distinct, and clearly separating lateral tubercles.

Sanderson: Species of Stenelmis 687

Basal tubercle rounded anteriorly, narrowed and elongate pos-
teriorly and usually distinctly carinate to near the pronotal base.
Anterior tubercle prominent and rounded.

Elytra. Length, 2.37-2.75 mm.; width, 1.25-1.4 mm. Each ely-
tron usually faintly bimaculate. Humeral spot on inside of sixth
interval, usually occupying no more than the fifth interval at base,
and extending along the interval only for about one fifth of its
length. Subapical spot extending from about one third behind the
middle of elytron to barely beyond end of carina of sixth interval.
First elytral stria complete from base to apex. Third interval
conspicuously elevated at base, the elevation ending posteriorly to
about the end of humeral spot. Elytral punctures deep on disk
and becoming finer posteriorly, never obsolete on apical declivity of
elytra.

Venter. Light brown to grey, usually lighter posteriorly. Apical
abdominal emargination less than width of last tarsal segment.

Legs. Hind tibiae, 1-1.15 mm.; middle tibiae, .8-.9mm. ; anterior
tibiae, .87-1 mm. Last tarsal segment distinctly longer than preced-
ing four combined. Granules of femora usually not evident. Femora
and tibiae of same general color as venter, the tarsi light brownish.

Notes on types. Holotype male, allotype female and two para-
types, Ireland, Miss., 7-8-34, R. H. Beamer. Additional paratypes
from Tallulah, La., VII-12-30, VII-13-30, and Vin-15-30, P. A.
Glick; Beauregard parish, Louisiana, 8-18-28, J. G. Shaw; Green
county, Mississippi, 3-3-1932, H. Dietrich, Gaines creek; Leaf,
Miss., Mar. 3, 1932, H. Dietrich; Lucedale, Miss., 6-22, P. N. Mus-
grave; Hope, Ark., VII-1-32, L. Knobel, at light (C. A. Frost
collection). The specimens from Tallulah, La., are from the Har-
vard College collection, that of C. A. Frost, and the Canadian na-
tional collection.

Remarks and comparative notes. This species, with S. douglasensis
n. sp., occupies a unique position in the Sinuata-hnmerosa group
by the complete absence of granules on the pronotum, legs, and
remainder of the body. The granules in this species and its ally
apparently are replaced by distinct punctures. S. grossa is dis-
tinguished from S. douglasensis by the bimaculate instead of vittate
elytra, and by the shape ,of the processes on the median lobe of the
aedeagus. In S. grossa the processes are subangulate anteriorly;
in *S. douglasensis the processes are evenly rounded. These are two
of our largest species of Stenelmis.

Location of types. Holotype, allotype, and paratypes in the

688 The University Science Bulletin

Francis Huntington Snow entomological collection, University of
Kansas. Additional paratypes in the Harvard College collection;
Canadian national collection; collection of Dr. Paul N. Musgrave,
Fairmont, W. Va.; and C. A. Frost, Framingham, Mass.

Stenelmis parva n. sp.

(PI. LXXX, fig. 14)

Size. Length, 2.5 mm. ; width, .87 mm.

Form and color. Body elongate, parallel, convex. Elytra entirely
immaculate above, color black.

Head. Median longitudinal band as wide as the two lateral ones
combined, the granulation between median band and eye irregular
and indistinct. Antennae and palpi yellow testaceous. Antennae
longer than pronotum.

Pronotum. Length, .75 mm.; width, .65 mm. Base of sulcus and
on either side of basal tubercles, light grey; remainder of pronotum
a dull, dark brown. Sides of pronotum very slightly rounded behind
middle, a little convergent posteriorly and anteriorly to basal and
apical angles; width at apex equal to width at base. Granules on
either side of sulcus very fine and separated by twice their diameters.
Median longitudinal sulcus distinct though not deep, the anterior
limits indistinct; sulcus extending posteriorly from about the apical
one third to near the base, the sides nearly parallel. Oblique im-
pression on either side of median sulcus distinct and clearly separat-
ing lateral tubercles. Basal tubercle a little elongate, the anterior
one fairly prominent and rounded.

Elytra. Length, 1.85 mm.; width, .87 mm. Elytra entirely im-
maculate, there being no trace of pale spots. First stria complete.
Third interval but slightly elevated at base. Elytral punctures
moderately large and deep on disk, never obsolete on declivity.

Venter. Blue-grey, the posterior margin of abdominal segments
and an entire narrow stripe on the last segment, lighter. Apical
emargination less than width of the last tarsal segment.

Legs. Hind tibiae, .75 mm. ; middle tibiae, .62 mm. ; anterior
tibiae, .65 mm. Last tarsal segment longer than preceding four
combined. Granules of femora separated by their own diameters.
Femora and tibiae darker grey than venter. Tarsi and extreme
apex of the tibiae reddish-brown.

Notes on types. Holotype male and one paratype, Latimer
county, Oklahoma, 12-VII-1931, W. Fisher. Allotype female, Edna,
Texas, 8-9-28, R. H. Beamer.

Sanderson: Species of Stenelmis 689

Remarks and comparative notes. S. parva n. sp. has a wider
median frontal band on the head than in any other species ex-
amined. This species is associated with S. antennalis n. sp. and
5. musgravei n. sp. by its small size, but is readily distinguished
by the total absence of elytral spots. By this characteristic it would
be placed with S. fuscata Blatch., from which it is separated by its
much smaller size.

Location of types. Holotype and paratypo in the collection of
Dr. Paul N. Musgrave. Allotype located in the Francis Huntington
Snow entomological collection, University of Kansas.

Stenebm's fuscata Blatchley

(PI. LXXX, fig. ].'))
1925. Stendmis fuscatiis Blatchley, Can. Ent. 57, p. 164.

Size. Length, 3.25-3.4 mm.; width, 1.15-1.25 mm.

For?n and Color. Body elongate, convex, parallel. Color of en-
tire upper surface a uniform dark brown without indication of spots
on vittae.

Head. Granulations between eye and frontal band rather coarse
and uneven. Antennae and palpi yellow-testaceous. Antennae
usually conspicuously shorter than pronotum.

Pronotum. Length, 1-1.1 mm.; width, .9-. 95 mm. Anterior angles
and margin of pronotum rufescent. Sides parallel on basal half,
sinuate before middle, then a little convergent and rounded to apex,
A slight sinuation between hind angles and rounded sides of pro-
notum. Width at base of pronotum equal to that at middle. Nar-
rower at apex than at base. Median sulcus moderately deep, di?
tinct to just before the base, and extending posteriorly from about
the apical one fourth. Oblique impression on each side of sulcus
distinct and clearly separating lateral tubercles. Basal tubercle a
little elongate, anterior one more rounded and conspicuous.

Elytra. Length, 2.37-2.5 mm.; width, 1.15-1.25 mm. Entirely
immaculate above with no trace of spots. First elytral stria com-
plete from base to apex. Third interval elevated at base. Elytral
punctures moderate on disk, then smaller toward apex, never ob-
solete.

Venter. Very dull grey to light brown, the abdominal segments
nearly uniform in color. Apical emargination about equal to width
of last tarsal segment.

Legs. Hind tibiae, 1 mm.; middle tibiae, .85 mm.; anterior tibiae,
.9 mm. Last tarsal segment longer than preceding four combined.

44_4141

690 The University Science Bulletin

Granules of femora rounded, separated by their own diameters or
less. Femora and tibiae of the same general color as venter. Tarsi
light brown.

Notes on types. The type and paratype, from Royal Palm Park,
Fla., 3-18-24, W. S. Blatchley, have been studied.

Remarks and comparative notes. The specimens of this species
before me are remarkably uniform in size. Stenelmis fuscata Blatch.
was the first truly immaculate species to be named from our fauna.
It is closely related to Stenelmis himgerfordi n. sp., from which it
may be separated by its comparatively large size, and by the very
distinct lateral processes on the median lobe of the aedeagus. In
S. hungerfordi these processes are only about one fourth the width
of the lateral lobes.

Notes on distribution. In addition to the types, the following
localities are represented in the material at hand: Fort Pierce, Fla.,
8-7-30, R. H. Beamer, P. W. Oman; Lacoochee, Fla., 8-18-30, J. 0.
Nottingham; Hillard, Fla., 8-19-30, P. W. Oman; and Hillard, Fla.,
8-31-30, J. 0. Nottingham.

Location of types. Type in the collection of Prof. W. S. Blatchley.
Paratype in the Francis Huntington Snow entomological collection,
University of Kansas.

Stenelmis hungerfordi n. sp.

(PI. LXXX, fig. 16)

Size. Length, 2.7-2.8 mm.; width, 1-1.1 mm.

Form and color. Body elongate, convex, parallel. Color of elytra
black, each elytron usually with pale humeral and subapical spots;
the humeral spot, if present, always on inside of sixth interval.

Head. Granulations between eye and band generally separated
by twice their diameters. Antennnae and palpi yellow-testaceous.
Antennae slightly longer than pronotum.

Pronotum. Length, .85-. 9 mm.; width, .75-. 8 mm. Color of pro-
notum dull grey to brown ; the anterior angles and margin rufescent.
Occasionally a little lighter betweeen basal tubercles. Sides nearly
parallel, though a little convergent of base; sinuate before the mid-
dle, then sides parallel, or slightly divergent at apices, never con-
vergent. Sides before the basal angles perceptibly sinuate. Width
at base of pronotum nearly equal to width behind middle, narrower
at apex than at base. Granules on either side of median sulcus
separated by about twice their diameters. Median longitudinal
sulcus moderately deep, convergent posteriorly, indistinct at ex-

Sanderson: Species of Stenelmis 691

treme base. Oblique impression on either side of median sulcus
distinct, clearly separating lateral tubercles. Basal tubercle a little
elongate, never carinate; anterior one a little less distinct and
rounded.

Elytra. Length, 1.95-2.1 nun.; width, 1-1.1 mm. Usually en-
tirely immaculate above, though occasionally a small, indistinct
humeral spot on inside of sixth interval, and a faint subapical one.
Spot, when present, covering no more than fifth interval. First
elytral stria complete from base to apex. Third interval elevated at
base. Elytral punctures large on disk, becoming rapidly smaller at
apex, never obsolete on apical declivity.

Venter. Dull grey to light brown, the abdominal segments gen-
erally lighter on posterior borders. Abdominal emargination less
than width of last tarsal segment.

Legs. Hind tibiae, .82-. 87 mm.; middle tibiae, .7-.72 mm.; an-
terior tibiae, .75 mm. Last tarsal segment longer than preceding
four combined. Granules of femora rounded and usually separated
by a little less than their own diameters. Upper surfaces of femora
and tibiae grey with lower surfaces light brown; tarsi a lighter
testaceous.

Notes on types. Holotype male, allotype female and numerous
paratypes, Lacooche, Fla., 8-18-30, J. 0. Nottingham, collected at
light. Additional paratypes from Ocala, Fla., 8-17-30, Paul W.
Oman; Plant City, Fla., 8-15-30, J. O. Nottingham.

Remarks and comparative notes. This species is named after Dr.
H. B. Hungerford, University of Kansas, who has aided me in many
ways in my study of Stenelmis. S. hungerfordi n. sp. is closely re-
lated to S. fuscata B latch., but is readily separated by its smaller
size, the antennae are longer than the pronotum, and an occasional
specimen shows a faint humeral and apical spot on each elytron.
The processes on the median lobe of aedeagus are evenly rounded
and no more than one fourth the width of the lateral lobe. In S.
juscata, the processes are fully one half the width of lateral lobe.
Reference to the plate of genitalia will show other minor differences.

Location of types. Holotype, allotype, and paratypes in the
Francis Huntington Snow entomological collection. University of
Kansas. Additional paratypes in the collection of Dr. Paul N.
Musgrave.

692 The University Science Bulletin

Stcnebnis hmnerosa Mots.

(PI. LXXX, fig. 17)

1859. Stenelmis humerosus Mots. Etudes Entomo. VIII, p. 50.
1869. Stenelmis linearis Zimm. Trans. Am. Ent. Soc. 11, p. 259.

Size. Length, 2.3-2.7 mm.; width, .95-1.1 mm.

Form and color. Body elongate, convex, nearly parallel. Color
of elytra dark brown to nearly black; each elytron with an entire
longitudinal vitta which may be somewhat separated on disk into a
humeral and a subapical spot. Vitta or spot covering humerus at
base.

Head. Granulations between eye and band rounded, separated by
several times their own diameters. Antennae and palpi light brown-
ish testaceous. Antennae equal in length to pronotum.

Pronotum. Length, .7-. 75 mm.; width, .68-.75 mm. Entire
median sulcus and from its anterior end to apex of pronotum fuscous.
On either side of sulcus a fuscous triangular area extending from
base of pronotum between sulcus and lateral tubercle to a point near
middle of pronotum. A small fuscous spot near anterior angles of
pronotum. Margins of sulcus and remainder of pronotum a dull
yellowish-grey. These areas with granules moderate in size though
nearly reduced to general surface, the granules separated by about
twice their own diameters. Sides of pronotum nearly parallel in
basal half, then slightly narrowed at middle and subparallel to apical
angles. Width at base of pronotum nearly equal to that at middle,
but slightly narrower at anterior angles than at base. Granules on
either side of median sulcus nearly indistinct. Median sulcus mod-
erate in depth, nearly obsolete on basal declivity and extending pos-
teriorly from near apical one third. Oblique impression on either
side of sulcus almost wanting and only perceptibly separating lat-
eral tubercles. Basal and anterior tubercles not at all conspicuous
and but little raised above the disk of pronotum.

Elytra. Length, 1.75-1.88 mm.; width, .95-1.1 mm. Each elytron
with the vitta usually entire, though somewhat clouded at middle.
Humeral part of vitta embracing umbone, covering fourth and fifth
elytral intervals and extending to a point about half way between
end of carina of sixth interval and elytral apex. First elytral stria
complete from base to apex. Third interval scarcely, if at all,
elevated at base. Elytral punctures deep on disk, then becoming
much smaller on apical declivity though never obsolete.

Venter. Grey or greenish-grey with nearly all of posterior half of
abdominal segments dull reddish-brown. Abdominal emargination
equal to width of last tarsal segment.

Sanderson: Species of Stenelmis 693

Legs. Hind tibiae, .75-.8 mm.; middle tibiae, .65-.72 mm.; an-
terior tibiae, .65-.72 mm. Last tarsal segment distinctly longer than
preceding four combined. Granules of femora generally rounded
and very closely placed, separated by less than their diameters.
Femora and tibiae entirely grey, the tarsi dark brown.

Notes on types. The type female of S. humerosa Mots., from
Tennessee, and the type male of S. linearis Zimm. from South
Carolina, have been examined. An examination shows them to be
the same species. In 1870 Horn expressed an opinion that the two
might be the same, although he had not seen the type of S. humerosa.
Since S. humerosa has priority by ten years, <S. linearis thereby be-
comes a synonym. The type of S. humerosa bears the following
label: "29 Juni vole a la mandelle dans la chambre dans uni nuit
organse. Alabam."

Remarks and comparative notes. The present species is closely
related to S. mirabilis n. sp., but is easily separated from the latter
by the testaceous instead of dark brown to black palpi. In addition
the tarsi are not lighter at base as described for S. mirabilis.

Notes on distribution. In addition to the types, four specimens
have been examined from the following localities: Md.: Offutts Isl.,
16.7.16, Loomis and Barber Coll. D. C: "D. C." Otto Lugger
Coll. Mass.: Chicopee, July 20, W. Knaus Coll.

Location of type. The female type of S. humerosa Mots, is located
in the Zoological Museum of Moscow, U. S. S. R. The type of S.
linearis Zimm. is in the LeConte collection at Harvard College.

Stenelmis mirabilis n. sp.

(PI. LXXX, fig. 18)

Size. Length, 2.7-2.9 mm.; width, 1.1-1.12 mm.

Form and color. Body elongate, convex, a little wider behind.
Elytra dark to black. Each elytron distinctly bimaculate and never
with the vitta entire. Humeral spot embracing umbone.

Head. Granulations between eye and band separated by about
twice their diameters. Antennae brownish-piceous to black, lighter
at base, the palpi piceous to black. Antennae slightly longer than
pronotum.

Pronotum. Length, .86 mm. ; width, .Id-.ll mm. Pronotum
usually dull grey to deep brown. Sides nearly parallel before the
base, narrowed anteriorly to middle, sinuate, then straight to apex.
A little narrower across the apex of pronotum than at base. Granules
on either side of median sulcus at base separated by about twice

694 The University Science Bulletin

their diameters. Median longitudinal sulcus distinct, moderately
deep, distinctly narrowed posteriorly, and extending from the apical
one fourth nearly to base. Oblique impression on either side of
median sulcus distinct and clearly separating lateral tubercles.
Basal tubercle rounded and but little elongate. Anterior tubercle
rounded, not prominent.

Elytra. Length, 1.95-2 mm.; width, 1.1-1.12 mm. Each elytron
distinctly bimaculate, the vitta never entire. Humeral spot em-
bracing the umbone, extending diagonally inside of the six-th inter-
val, and usually covering a little more than the fifth interval for
one fourth or one fifth of its length. Subapical spot about equal to
the dark band separating it from the humeral spot. Subapical spot
extending to a point a little beyond end of carina of sixth interval.
First elytral stria complete from base to apex. Punctures moder-
ately deep on disk, becoming smaller and very fine on declivity,
though never entirely absent.

Venter. Dull brown to a blue-grey. Apical abdominal emargina-
tion about equal to the width of last tarsal segment.

Legs. Hind tibiae, .8-. 85 mm.; middle tibiae, .62 mm.; anterior
tibiae, .65 mm. Last tarsal segment distinctly longer than preceding
four combined. Granules on hind femora separated by no more
than their own diameter. Femora above grey, usually a light brown
below at base; tibiae nearly entirely testaceous except extreme base
and apices, which are colored as the femora. Tarsi dark brown to
black, the basal two thirds of the last segment lighter.

Notes on tijpes. Holotype male, allotype female, Cos. Cob. Ct.
(H. H. Knight Coll.) ; one paratype, Cos. Cob. Ct. (W. Knaus Coll.) ;
one paratype, "N. C." (Otto Lugger Coll.) ; and an additional para-
type from Anderson, S. C. Jl. S, P. N. Musgrave (P. N. Musgrave
Coll.).

Remarks and comparative notes. This species is of nearly the
same size and form of S. humerosa Mots., but is easily distinguished
by its lighter tibiae and much darker tarsi. In S. humerosa the tibiae
are of the same color as the femora. The cleaner examples of this
species are very striking in their type of coloration of the legs and
elytra.

Location of types. Holotype and allotype located in the Francis
Huntington Snow entomological collection, University of Kansas.
Paratypes in the W. Knaus collection now located at Kansas State
College, Manhattan, P. N. Musgrave collection, and that of the Uni-
versity of Minnesota.

Sanderson: Species of Stenelmis 695

Stenelmis antennalis n. sp.

(PI. LXXX, fig. 19)

Size. Length, 2.5-2.7 mm. ; width, 1 mm.

Forin and color. Body elongate, parallel, convex. Color of elytra
deep brown to black. Each elytron with humeral and subapical
testaceous or reddish spots which are always on the inside of the
sixth interval.

Head. Antennae with the last six or seven segments nearly always
black, the basal segments testaceous. Palpi testaceous. Antennae
longer than pronotum.

Pronotum. Length, .75-. 85 mm.; width, .65-.7 mm. Basal one
third of pronotum and margins of median sulcus often dull grey, the
remainder dull brown to black. Anterior angles and margin faintly
rufescent. Sides rounded behind middle, a little convergent posteri-
orly; sinuate before the middle, then sides parallel to apex. Width
at base of pronotum ecjual to width at apex. Granules on either side
of median sulcus very fine and generally separated from two to
several times their diameters. Median longitudinal sulcus very
shallow, a little narrowed behind and extending from the apical one
third or one fourth to near the base. Oblique impression on either
side of median sulcus often indistinct and faintly separating lateral
tubercles. Basal tubercle faintly raised and elongate, the anterior
one but scarcely evident.

Elytra. Length, 1.85-1.95 mm.; width, 1 mm. Each elytron with
a pale or reddish humeral spot and a subapical one which extends
but little beyond apex of carina of sixth interval. Spots covering
most of the fourth and fifth intervals, and never uniting to form an
entire stripe. First elytral stria complete. Third interval only
faintly elevated at base. Elytral punctures rather shallow on disk,
moderate in size, and usually extending to apex, though becoming
much finer and occasionally obsolete on declivity.

Venter. Dull brown to grey, the posterior margin of abdominal
segments usually lighter. Apical emargination equal to width of last
tarsal segment.

Legs. Hind tibiae, .75 mm. ; middle tibiae, .6-.62 mm. ; anterior
tibiae, .62-. 64 mm. Last tarsal segment longer than preceding four
combined. Granules of femora separated by a little less than their
own diameters. Femora of same general color as venter. Tibiae
and tarsi a little lighter.

Notes on types. Holotype male, Lucedale, Miss., Mar. 1, 1932, H.
Dietrich (P. N. Musgrave Coll.) ; allotype female, Hilliard, Fla.,

696 The University Science Bulletin

8-31-30, J. 0. Nottingham; additional paratypes from the following
localities: Mobile, Ala., VI 6, 27, Darlington; DeFuniak Springs,
Fla., June 28, 31, P. N. Musgrave; Whiskey creek. Miss., Leaf III
4, 1932, H. Dietrich; Lucedale, Miss., June 22-23, Cedar creek, P. N.
Musgrave; Lucedale, Miss., 7-5-31, H. Dietrich; Clara, Miss., Wayne
county, June 26, 31.

Remarks and comparative notes. The uniform lighter palpi and
darker antennae, comparatively narrower pronotum and less ele-
vated third elytral interval will easily distinguish this species from
■S. musgravei n. sp., to which it is related.

Location of types. Holotype and paratypes in the collection of
Dr. Paul N. Musgrave. Allotype and paratypes in the Francis
Huntington Snow entomological collection. University of Kansas.
.Additional paratypes in the Harvard College collection.

Stenelmis quadrimaculata Horn

(PI. LXXXI, fig. 2)

1870. Stenelmis Quadrimaculatus Horn, Trans. Am. Ent. Soc. Ill, p. 40.

1933. Steyielmis blatchleyi Musgrave, Proc. Ent. Soc. Wash., Vol. 35, No. 4, p. .57.

1910. 11 Stenelmis sulcatus Blatchley, Col. of Ind. p. 681.

Size. Length, 2.7-3.2 mm.; width, 1.1-1.25 mm.

Form and color. Body elongate, convex, nearly parallel. Each
elytron usually distinctly bimaculate and never with the vitta entire.
Humeral spot on inside of sixth interval.

Head. Granulations between eye and band very fine, rounded,
and separated by several times their own diameters. Antennae
brownish, nearly piceous in some examples, but always lighter at
base. The palpi deep brown to piceous. Antennae as long as
pronotum.

Pronotum. Length, .86-1 mm.; width, .77-. 87 mm. Inside of
sulcus from near its base and occasionally a small area on either side,
fuscous. Remainder of pronotum dull yellowish-grey. Sides nearly
parallel in basal half, then gradually rounded and subparallel to
apex. Slightly wider across rounded sides of pronotum than at base.
Apex distinctly shorter than base. Granules on either side of sul-
cus at base separated by about twice their own diameters. Median
longitudinal sulcus distinct, slightly narrowed posteriorly, and ex-
tending from about apical one fourth to base. Oblique impression on
either side of sulcus distinct and clearly separating lateral tubercles.
Basal tubercle rounded.

Elytra. Length, 2-2.35 mm.; width, 1.1-1.25 mm. Each elytron
distinctly bimaculate, with the vitta never entire. Humeral spot on
inside of sixth interval and occupying nearly all of the fourth and

Sanderson: Species of Stenelmis 697

fifth intervals. Band separating vittae jiist slightly shorter than
humeral spot. Subapical spot extending a little beyond apex of
carina of sixth interval. First elytral stria complete from base to
apex. Tiiird interval distinctly though not conspicuously elevated
near base. Elytral punctures deep on disk, then becoming smaller
toward apex of elytra, but never obsolete on declivity.

Venter. Dull red or brownish to greenish-grey with abdominal
segments generally dull brown on posterior half. Apical emargina-
tion about same as width of last tarsal segment.

Legs. Hind tibiae, .85-.9 mm.; middle tibiae, .65-.75 mm.; an-
terior tibiae, .7-.82 mm. Last tarsal segment distinctly longer than
preceding four combined. Granules of femora occasionally sep-
arated by as much as their own diameters, though generally closely
placed. Femora and tibiae of the same color as venter, though the
tibiae sometimes faintly lighter. Tarsi reddish-brown.

Notes on types. Cotypes of S. quadrimaculata Horn, from Ben-
nington county, Vt., and the type of S. sulcata Blatch., from Marshall
county, Ind., have been examined. They are apparently the same.
Variation of pronotal sculpture within many of our species might
account for the present species being known under two names.
Since *S. quadrimaculata has priority by forty years, S. sidcata
should be dropped from our list. S. blatchleyi Musgrave was the
new name proposed for S. sulcata Blatchley, since the latter name
was preoccupied by S. sulcata Grouvelle, a species described from
Sumatra in 1892 (Not. Leyd. Mus. XIV, p. 188).

Remarks and comparative notes. S. quadrimaculata Horn is very
closely related to S. musgravei n. sp. It may be distinguished by its
larger size, less extensive fuscous areas on pronotum, and by certain
features of the genitalia. In both species the median lobe of aedeagus
possesses evenly rounded lateral processes. In S. quadrimaculata
these processes are broader than in S. musgravei, being about two
thirds the width of the median lobe. The median lobe is more
pointed and the extreme bases of the lateral lobes are more evenly
rounded. In S. musgravei, the tip of median lobe is evenly rounded,
and the bases of lateral lobes nearly subangulate externally.

Notes on distribution. In addition to the types from Vermont,
material from the following localities has been examined:

United States:

Indiana: Marshall county, 6-23-34, M. W. Sanderson.
Michigan: Leelenau county, 8-15-33, H. T. Peters: Livingston county,
. June-July, H. B. Hungerford, Bass L.; Cheboygan county, July,
1936, M. W. Sanderson, Black Lake.

698 The University Science Bulletin

New York: Buffalo (F. C. Bowditch Coll. at Harvard College) ; Buffalo

(Otto Luggar collection at Univ. of Minn.).
District of Columbia: Washington (H. H. Knight collection).

Canad.a :

Canada: "Can." (H. H. Knight collection).

Quebec : Wakefield, 20. VII 1932, W. J. Brown (Canadian national col-
lection); Knowlton's Landing, 10, VII 1927, W. J. Brown (Canadian
national collection).

Location of types. Cotypes in the Horn collection at the Phila-
delphia Academy of Sciences. The type of S. sulcata Blatch. is in
the collection of Prof. W. S. Blatchley, Indianapolis, Ind.

Stenelmis musgravei n. sp.

(PI. LXXXI, fig. 3)

Size. Length, 2.4-2.65 mm.; width, .95-1.05 mm.

Form and color. Body elongate, convex, nearly parallel. Elytra
black, each elytron usually distinctly bimaculate and at least a
median clouding on vitta. Humeral spot usually on the inside of the
sixth interval.

Head. Granulations between eye and band separated from two
to three times their diameters. Antennae and palpi dusky to piceous,
the palpi ordinarily darker. Antennae about equal in length to
pronotum.

Pronotum. Length, .77-. 8 mm.; width, .67 mm. Entire median
sulcus to apex of pronotum, and an elongate triangular area on
either side of sulcus to a point about two thirds from base — deep
brown to black. Remainder of pronotum dull yellowish-grey. Sides
of pronotum a little convergent from rounded sides to base. Nar-
rowed about the middle, then subparallel to apex. Slightly narrower
at apex of pronotum than at base. Granules on either side of
median sulcus separated by once or twice their own diameters.
Median longitudinal sulcus distinct, narrowed posteriorly, and ex-
tending from about the apical one fourth or one fifth to near base.
Oblique impression on either side of median sulcus moderately deep
and separating lateral tubercles. Basal tubercle rounded, a little
elongate; the anterior tubercle rounded, not prominent.

Ehjtra. Length, 1.75-1.85 mm.; width, .95-1.05 mm. Each ely-
tron nearly always distinctly bimaculate, and with the humeral spot
usually on the inside of the sixth interval. Maculae occupying nearly
all of fourth and fifth intervals according to limitations of spot.
Band separating spots variable in size, but never entirely absent,
occasionally reduced to a faint cloud. Subapical spot terminating

{

Sanderson: Species of Stenelmis 699

beyond apex of carina of sixth interval. First stria complete from
base to apex. Third interval distinctly elevated at base. Elytral
punctures deep on disk, finer posteriorly, never obsolete on declivity.

Venter. Light brown to a greenish-grey, the posterior margin
and median longitudinal band on last segment lighter. Apical
emargination equal to width of last tarsal segment.

Legs. Hind tibiae, .75 mm.; middle tibiae, .62-. 65 mm.; anterior
tibiae, .64-. 67 mm. Last tarsal segment distinctly longer than pre-
ceding four combined. Granules of femora separated by less than
their own diameters. Femora and tibiae of the same color as venter.
T.arsi dark brown to black, the basal two thirds of last segment often
a little lighter.

Notes on types. Holotype male, allotype female, and one para-
type, Winchester, Va., 7-21-34, M. W. Sanderson. Additional para-
types from Schellburg, Pa., VIII 16; Wardenville, W. Va., July 19,
P. N. Musgrave, Trout Run; Sleepy creek. Route 9, Morgan county,
W. Va., July 23, 1933; Offutts Isl, Md., 16-7-16, Loomis and Barber,
at light; Hollister, Mo., 7-28-34, M. W. Sanderson; and Batavia,
N. Y., Sept., 1915, H. H. Knight.

Remarks and comparative notes. As indicated after the descrip-
tion of S. quadrimaculata Horn, this species is closely related, but is
readily separated by its smaller size. The genitalia are of the same
general type but exhibit the differences indicated under >S. quadri-
maculata Horn. It shows some affinities with S. antennalis n. sp., but
is easily separated by the color of the palpi, which are always as dark
as or darker than the antennae. The palpi are always testaceous or
lighter in S. antennalis n. sp. In two examples of S. musgravei n. sp.
the humeral spot covers a part of the umbone. However, this is not
usual and is not to be regarded as characteristic of the species. These
specimens, in this particular, come close to S. humerosa Mots., but
are distinguished by the distinctly raised third intervals.

Location of types. Holotype, allotype, and paratypes in the
Francis Huntington Snow entomological collection. University of
Kansas. Additional paratypes in the collection of Dr. Paul N.
Musgrave.

Stenelmis sinuata Lee.

1802. Stenelmis sirntatus Lee. Proc. Ac. N. S. Phila. VI, p. 44.

Size. Length, 3.25-3.45 mm.; width, 1.2-1.35 mm.

Form and color. Body elongate, moderately subdepressed, dis-
tinctly inflated behind. Elytra shiny brown; each elytron with a
faint longitudinal vitta on inside of sixth interval.

700 The University Science Bulletin

Head. Granulations between eye and band small but prominent
and separated from two to four times their diameters. Antennae
and palpi testaceous. Antennae considerably shorter than pro-
notum.

Pronotum. Length, 1-1.1 mm.; width, .87-. 95 mm. Disk of
pronotum light brown, but faintly shining. Pronotum generally
widest behind middle, then a little convergent to the hind angles, be-
fore which there is frequently a slight sinuation. Sides rather sud-
denly narrowed before the middle, deeply sinuate, the anterior
angles usually strongly divergent. The extreme apical angle
truncate, the truncation sometimes obtuse. Pronotum usually as
wide at base as behind the middle, but never wider at base. Width
across the anterior angles but little less than the width at base.
Granules on either side of median sulcus at base separated by about
twice their diameters. Median longitudinal sulcus very distinctly
narrowed posteriorly from near the apical one fourth to the base.
Oblique impression on either side of median sulcus distinct, clearly
separating lateral tubercles. Basal tubercle elongate, but not dis-
tinctly carinate behind. Anterior tubercle moderately elevated and
rounded.

Elytra. Length, 2.35-2.5 mm.; width, 1.2-1.35 mm. Each elytron
maculate. The vitta only narrow and occupying but little more
than fifth interval. First elytral stria complete from base to apex.
Third interval conspicuously and acutely elevated at base. The
elevation occupying no more than one fourth of the total length of
the interval. Elytral punctures are deep on disc and very conspicu-
ous to the apex.

Venter. Nearly uniformly dark brown in the South Carolina
specimen. In all others the color is not distinguishable, due to a fine
coat of foreign material. Emargination of last abdominal segment
a little less than width of last tarsal segment.

Notes on type. The type is a female and is labeled Tallulah, Ga.

Remarks and comparative notes. No males of this species have
been examined, though eleven specimens are before me. Most of the
examples are covered with vegetable debris, and in some individuals
the vittae are all but invisible. This species perhaps shows its
closest relationship to 5. vittipennis Zimm. and allies, but is readily
separated by having the sides of pronotum strongly divergent in
apical third. The apical angles of pronotum are not acute, as in
most of our species, but subtruncate. It is nearly the size of S.
grossa n. sp. and S. douglasensis n. sp., but differs by its granulate
instead of punctate legs, pronotum, and venter.

Sanderson : Species of Stenelmis 701

Notes on distribution. In addition to the type from Tallulah, Ga.,

the following localities are represented in the material at hand:

Georgia: Wrens, July 7, 1931, P. N. Miisgrave.
Alabama: Mobile, June 25, 1931, P. N. Musgrave.
South Carolina: "S. C." (Harvard College collection).
Mississippi: Lucedale, June, P. N. Musgrave, in Reedy creek.

Location of type. Type in the LeConte collection at Harvard
College.

Stenelmis decorata n. sp.

(PL LXXXI, fig. 4)

Size. Length, 2.87-3 mm.; width, 1.1-1.15 mm.

Form and color. Body elongate, usually distinctly wider behind,
convex. Each elytron with an entire yellowish vitta which is often
clouded medially, though not distinctly bimaculate. Vitta on inside
of sixth interval.

Head. Granules between eye and band separated from two to
three times their own diameters. Antennae and palpi yellow-testace-
ous, the antennae a little longer than pronotum.

Pronotiim. Length, .88-. 95 mm.; width, .75-.8 mm.' Color vary-
ing from deep brown to light grey, basal one third or more, often
lighter. Sides rounded just behind the middle, then almost parallel
to base, scarcely sinuate before the hind angles. Narrowed and
sinuate before the rounded sides, then parallel to apex. Width of
pronotum at apex but little less than width at base. Granules im-
mediately on either side of median sulcus at base separated by twice
their diameters. Median longitudinal sulcus deep, extending from
near apical one fourth of pronotum to base, often a little narrowed
posteriorly. Oblique impression distinct and clearly separating
lateral tubercles. Basal tubercle small, slightly elongate though
never extending to pronotal base ; anterior tubercle rounded, distinct
but not prominent.

Elytra. Length, 2.12-2.32 mm. ; width, 1.1-1.15 mm. Each elytron
with the vitta entire, on the inside of sixth interval, covering the
fourth and fifth intervals, and extending a little beyond apex of
carina of sixth. First elytral stria complete from base to apex.
Third interval elevated on basal declivity though usually not acute.
Elytral punctures on disk moderate in depth and size, smaller toward
apex, never obsolete on declivity.

Venter. Light brown to grey, posterior borders of abdominal seg-
ments ordinarily lighter. Apical emargination equal to width of last
tarsal segment.

702 The University Science Bulletin

Legs. Hind tibiae, .9-1.1 mm.; middle tibiae, .75-. 85 mm.; an-
terior tibiae, .8-. 9 mm. Last tarsal segment a little longer than four
preceding combined. Femora and tibiae uniformly dark brown to
grey, the tarsi entirely testaceous.

Notes on types. Holotype male, allotype female, Lawrence, Kan.,
8-10-32, M. W. Sanderson, at light. Paratypes collected from Juno
28-Sept. 24, 1930-1932, Lawrence, Kan., at light. Additional para-
types from Douglas county, Kansas, F. H. Snow; Washington, D. C.
(H. H. Knight Coll.); "Ind." (Otto Lugger Coll.); Pike county,
111., 7-10-34, Lowry and Hack.

Remarks and coniparative notes. Comparison of the hind tarsi
of S. decorata n. sp. with those of S. rittipennis Zimm. and S.
mdrkelii Mots, shows that those of the present species are relatively
longer and more slender. The pronotum is usually a little narrower,
and the whole aspect is a more slender one than the other two species.
It is also on the average smaller. It is readily distinguished from S.
hunger jordi n. sp., with which it is associated by genital characters,
by its narrower form, and distinctly maculate elytra. It is separated
from »S. vittipennis Zimm. and S. mdrkelii Mots, by the absence of
lateral processes on the median lobe of aedeagus.

Location of types. Holotype, allotype, and paratypes in the
Francis Huntington Snow entomological collection, University of
Kansas. Additional paratypes in the collection of Dr. Paul N.
Musgrave.

Stenelmis vittipennis Zimm.

(PI. LXXXI, figs. 5, 9, 10, 12)
1869. Stenelmis vittiperiDis Zimm. Trans. Am. Ent. Soc. II, p. 259.

Size. Length, 3-3.4 mm.; width, 1.1-1.35 mm.

Form and color. Body elongate, moderately convex, nearly paral-
lel. Color of elytra very dark brown to black. Each elytron with
an entire testaceous vitta confined to inside of sixth interval.

Head. Granules between eye and band minute, usually sep-
arated by several times their diameters, in some cases nearly absent.
Antennae and palpi yellow-testaceous. Antennae equal to or
slightly shorter than pronotum.

Pronotum. Length, .87-1.05 mm.; width, .8-. 95 mm. Color, dull
grey, the sulcus fuscous anteriorly. Basal one third or more of sides
subparallel. Margin narrowed anteriorly, sinuate and with sides
straight to apical angles. Granules on either side of median sulcus
at base minute, separated from two to several times their own
diameters. Median longitudinal sulcus usually moderate in depth

Sanderson : Species of Stenelmis 703

with sides a little narrowed posteriorly; extending from apical one
fourth nearly to base. Oblique impression on either side of sulcus
separating lateral tubercles, the impression usually faint. Basal
tubercle but slightly raised above the disk and a little elongate,
never carinate; anterior tubercle usually rounded and inconspicuous.

Elytra. Length, 2.25-2.6 mm.; width, 1.1-1.35 mm. Each elytron
with vitta entire, on inside of sixth interval, and extending posteri-
orly to a point usually half way between apex of elytron and carina
of sixth interval. Vitta covering fourth and fifth intervals, faintly
clouded medially in some specimens. First interval elevated at base,
though occasionally but little raised. Elytral punctures moderate
on disk; finer toward apex, but never obsolete.

Venter. Very light greyish-blue to darker grey, the abdomen dis-
tinctly iridescent. The last segment with apical margin usually
lighter and which continues as a narrow band to base. Apical ab-
dominal emargination equal to width of tarsal segment.

Legs. Hind tibiae, .97-1.12 mm.; middle tibiae, .75-. 87 mm.; an-
terior tibiae, .87-. 97 mm. Last tarsal segment distinctly longer than
four preceding combined. Granules on femora very small, rounded,
and separated from about two to four times their diameters. Tibiae
and tarsi uniformly colored as venter, though less iridescent.

Notes on type. The type of this species from South Carolina has
been examined.

Remarks and comparative notes. The present species is closely
related to S. mdrkelii Mots, and S. decorata n. sp. It may be sep-
arated from these two species by the subangulate lateral processes of
the median lobe of aedeagus. It also averages a little larger in size
than either of the other two species.

Notes on distribution. In addition to tlie type, the species has
been examined from the following localities:
United St.^^tes:

Kansas: Lawrence (Douglas Co.), June 28-Aug. 25, 1930-1932, M. W.
Sanderson and L. S. Henderson; Kiowa county, July 5, 1932, R. H.
Beamer; Riley county, July 12-19, J. B. Norton (Kan. State College
Coll.) ; Greenwood county, 8-1-1923, Beamer and Lawson.

New Jersey: Spotwood (H. H. Knight collection).

West Virginia: Fairmont, Aug. 3, P. N. Musgrave, at light.

Canada :

Quebec: Wakefield, 20-VII 1932, W. J. Brown (Canadian national collec-
tion) ; Knowlton, 29. VII. 1930, L. J. Milne.

Manitoba: Aweme, on various dates as follows: 28-30 VIII, 1917. N.
Criddle; 13 VIII, 1932, N. Criddle; 12 IX 1922, P. M. Whitehall
from Canadian national collection).

704 The University Science Bulletin

Location of type. Type in the LeConte collection, Harvard Col-
lege.

Stenelmis convexula n. sp.

(PI. LXXXI, fig. 8)

Size. Length, 2.75-3.1 mm. ; width, 1.1-1.12 mm.

Form and color. Elytra reddish brown to dull grey. Body
elongate, convex, distinctly wider behind. Each elytron with a faint
humeral and subapical spot. Humeral spot on inside of sixth in-
terval.

Head. Granulation between eye and band separated by twice
their own diameters. Antennae and palpi testaceous. Antennae
about ec^ual to length of pronotum.

Pronotum. Length, .87-. 9 mm.; width, .75-. 85 mm. Disk of
pronotum dull brown to grey. Anterior margin and angles faintly
rufescent. Pronotum widest behind the middle. Slightly convergent
to base and generally sinuate before the hind angles ; narrowed be-
fore the rounded sides, then sides parallel to apex. Pronotum wider
at base than apex. Granulations on either side of median sulcus very
fine, separated by about three times their own diameters. Median
longitudinal sulcus shallow and generally occupying only the median
one third of pronotum. Oblique impression on either side ill-defined
and barely separating lateral tubercles. Posterior tubercle rounded,
not at all elongate; the anterior tubercle nearly indistinct.

Elytra. Length, 2-2.25 mm.; width, 1.1-1.12 mm. Each elytron
usually with very faint humeral and subapical spots. Spots on in-
side of sixth interval and occupying no more than fifth interval at
base, subapical spot a little broader and terminating about the end
of carina of sixth interval. First elytral stria complete. Third in-
terval but very slight and indistinctly elevated at base. Elytral
punctures moderate on disk, becoming finer toward apex and in some
examples nearly obsolete on declivity.

Venter. Dull brown to grey. The abdominal segments a little
lighter medially. Last ventral segment with a narrow apical emargi-
nation equal to width of last tarsal segment.

Legs. Hind tibiae, .75-. 87 mm.; middle tibiae, .62-. 7 mm.; an-
terior tibiae, .65-.75 mm. Last tarsal segment longer than preceding
four combined. Granules of femur separated by no more than their
own diameters. Femora and tibiae colored above as venter. Femora
noticeably lighter below. Apex of femur, base and apex of tibia
lighter. Tarsi reddish brown.

Sanderson : Species of Stenelmis 705

Notes on types. Holotypc male, allotype female and twenty-five
paratypes, DeFuniak Springs, Fla., June 28, 1931, P. N. Musgrave.

Remarks and comparative notes. This species bears some re-
semblance to <S. juscata Blatch. and S. hungerfordi n. sp. It differs
from the former by the possession of elytral spots, and from this
species and S. hungerfordi, it differs by its more convex and broader
form. The lateral processes of aedeagus are rounded but broader
than 5. mdrkelii Mots. 8. convexula n. sp. appears to be a little
more convex through the humeri than other species in our fauna.

Location of types. Holotype, allotype, and paratypes in the col-
lection of Dr. Paul N. Musgrave, Fairmont, W. Va. Additional
paratypes in the Francis Huntington Snow entomological collec-
tion, University of Kansas.

Stenelmis mdrkelii Mots.

(PI. LXXXI, figs. 11, 14, 16)

1854. Stenelmis mdrkelii Mots. Etudes Entomologiques, p. 12.
1859. Stenelmis mdrkelii Mots. Etudes Entomologiques, p. 50.

Size. Length, 3-3.25 mm.; width, 1.17-1.3 mm.

Form and color. Body elongate, convex, nearly parallel. Color
of elytra piceous-brown to black, each elytron with an entire longi-
tudinal testaceous vitta which is confined to inside of sixth interval".

Head. Gi'anules between eye and band separated from one to
several times their own diameter. Antennae and palpi light brown-
ish-testaceous. Antennae about equal in length to pronotum.

Pronotum. Length, .9-1 mm.; width, .85-. 9 mm. Median sulcus,
and from its anterior extremity to apex of pronotum, fuscous. Re-
mainder of pronotum dull grey. Sides rounded behind the middle,
then slightly convergent to base. Narrowed anteriorly, with a
shallow sinuation before anterior angles. Granules on either side
of sulcus at base, and entire upper surface, separated by about twice
their diameters. Median longitudinal sulcus moderate in depth,
narrowed posteriorly and extending from apical one third to near
the base. Oblique impression on either side of median sulcus in-
conspicuous and shallowly separating lateral tubercles; the tuber-
cles rounded and barely evident.

Elytra. Length, 2.25-2.4 mm.; width, 1.17-1.3 mm. Each elytron
with the vitta entire, on inside of sixth interval, and extending
posteriorly to a little beyond apex of carina of sixth interval. Vitta
covering fourth and fifth intervals except at middle, where vitta is
slightly narrowed. First elytral stria complete from base to apex.
Third interval in type just perceptibly elevated near base. Elytral
45-4141

706 The University Science Bulletin

punctures moderate in size and depth on disk, never obsolete on
apical declivity.

Venter. Light grey to brown with the abdominal segments nearly
uniform in coloration. Apical abdominal emargination equal to
width of last tarsal segment.

Legs. Hind tibiae, .9-1.12 mm.; middle tibiae, .8-. 9 mm; anterior
tibiae, .85-1 mm. Last tarsal segment distinctly longer than pre-
ceding four segments combined. Granules on femora of most ex-
amples generally separated from once to twice their diameters.
Femora and tibiae entirely grey, the tarsi reddish-brown.

Notes on types. The type series of four males bearing the label:
"Stenelmis Miirkelii Motsch. Teness." has been examined. One of
these specimens has been dissected and designated as lectotype.
The granules on the femora of three of the specimens are a little
more closely placed than on the specimen selected as the type.

Remarks and comparative notes. This species is very closely re-
lated to S. vittipennis Zimm. and is only satisfactorily separated
from that species by the character of the aedeagus. The processes
of the median lobe of S. mdrkelii Mots, are evenly rounded, and not
subangulate anteriorly, as in S. vittipennis Zimm. There is also a
close resemblance to S. decorata n. sp., but the latter has the median
lobe without processes.

Notes on distribution. In addition to the type series from Ten-
nessee, the species has been examined from:

Arkansas: Berryville, 7-4-34, R. H. Beamer, at light.
Missouri: Hollister, 7-28-34, M. W. Sanderson, Long Creek.
Massachusetts: Chicopee (W. Knaus collection).
Pennsylvania: "Penn." (Otto Lugger collection) ; "Pa."

Location of types. In the Zoological Museum of Moscow, U. S.
S. R.

ADDENDA

The following two species of Stenelmis are the only known West-
ern Hemisphere species not found in the United States. Although
neither is closely related to the others, their original descriptions
are added here for the sake of completeness.

Stenelmis geayi Grouv.

1908. Stenelmis Geayi Grouv. Bull, du Mus. d'Hist. Nat., 14, pp. 181-182.

ORIGINAL DESCRIPTION

"Oblongo-elongata, subparallela, convexa, opaca, nigrofumosa,
maculis griseis in capite prothoraceque, avellaneis in elytris varie-
gatus. Antennae tenues, rufo-testacese, articulis omnibus multo

Sanderson : Species of Stenelmis 707

longioribus quam latis. Caput transverso-oblongum, margine antico
ante bases antennariim inflexo; oculis magnis, satis admotis, labro
rufo-testaceo. Prothorax antice valde angustus vix longior quam
ad basin latus, in longitudinem subsulacatus; baud dense punctu-
latus, punctis ad latera majoribus; margine antico valde producto,
medio et utrinque sinuato; lateribus arcuatis, ad basin sinuatis;
basi trisinuata; angulie anticis late obtusis, subhebetatis, posticis
acutis, extus productis. Scutellum suborbiculare. Elytra pro-
thorace latiora, ad apiceum subseparatim acuminata, magis duplo
longiora quam simul lata, punctato-striata; intervallis striarum
punctis latioribus; humeris calosis, subdentatis; disco ad apiceum
sat abrupte declivo, intervallo 6° ad initium partis inclinatae per
tuberculium acuminatum armato. Tarsi rufo-testacei. Long., 5-5
millim. 5.

"Oblong, allonge, subparallele, convexe, moir, grisatre, legerement
violace, varie de taches grises sur la tete et le prothorax; grises,
legerement rosees sur les elytres; taches mal limitees, formant
j)rincipalement sur le prothorax une band anterieure mediane et
deux bandes laterales, basilaires, et sur les elytres une serie de
bandes lineaires couvrant les stries, se soudant parfois et groupees
a la base sur les 2', 3\ 4" et 5^ stries, sur les cotes sur les 4%
5% et 6*, sur le disque vers la partie declive, sur les 2% et 3^; strie
sutturale presque entierement couverte par une tache lineaire, sou-
vent interrompue sur les intervalles de points. Antennes roux-
testace greles ; tous les articles beaucoup plus dongs que larges. Tete
en ovale transversal, inflechie un peu en avant de la naissance des
antennes, cellesci relativement rapprochees a la base; yeus gros;
labre roux testace; dernier article des palpes maxillaires allonge,
oblong. Prothorax fortement retreci en avant, un peu plus long que
large a la base, longitudinalement subsillonne dans le milieu, cou-
vert d'une punctuation profonds, peu serree, fine sur le disque, plus
forte sur les cotes; bord anterieur fortement saillant en avant,
arrondi, sinue dans le milieu et de chaque cote; bords lateiau
firrondis, sinues avant la base; base trisinuee; angles anterieurs
largement obtus, posterieurs aigus, reflechis en dehors; sur le disque
de chaque cote du sillon longitudinal, vers le premier tiers a partir
de la base, une petite elevation conique. Ecusson suborbiculaire.
Elytres plus larges que le prothorax, subacuminee separement au
sommet, environ deux fois et un tiers aussi longs que larges en-
semble, ponctues-stries; intervalles des stries plus larges que les
points, ceux-ci plus forts a la base qu'ar sommet; partie apicale des
elytres assez fortement et brusquement declive; calus humeraux

708 The University Science Bulletin

allonges, subdentes; 8* intervalle externe legerement carene, termine
vers la partie declive de I'elytre par un tubercule acumine. Tarses
rouxtestace. Metasternum fortement creuse pour recevoir la saillie
du prosternum ; saillie du premier segment ab dominal tres largement
obtuse.

"II exemplaires. — Guyane; riviere Lunier (F. Geay). Collection
du Museum de Paris.

"Espece remarquable par les taches d'un aspect limoneux, mal
definies qui decorent son prothorax et ses elytres."

Notes

Two specimens of this species from the type series have been
studied. It is not closely related to any species found within the
United States although it possesses certain characters peculiar to
the Simiata-humerosa group. The last tarsal segment is very much
longer than the four preceding combined, and the claw is propor-
tionately larger. The punctures of the pronotum are very fine, a
character which is found only in S. grossa Sand, and S. douglasensis
Sand, both of which are in the Sinuata-humerosa group. Other-
wise *S. geayi Grouv. departs from all other species in the Western
Hemisphere in many characters. The entire body is peculiarly
patterned with dark and light grey which extends on to the legs;
the antennae are very slender and are longer than the pronotum
by the last three segments. The antennae and palpi are brownish-
testaceous. The pronotal sulcus is shallow and extends nearly the
full length of pronotum. The margin of pronotum at the anterior
end of sulcus is distinctly emarginate. On either side of sulcus, near
basal third, is a rounded prominence, and opposite this, near lateral
margin, is another slightly smaller prominence. The third elytral
interval is elevated a little behind extreme base and with the in-
tervals on either side slightly prominent. The sixth interval is
elevated and prominent at extreme base. In the apical fourth of
the seventh elytral interval, there is a conspicuous dentiform process.
The apices of the elytra are separate.

For the present I must regard this as congeneric with the European
Stenelmis canaliculata (Gyll.) although its facies and robustness are
quite unlike other species known to me. In general appearance it
recalls certain Curculionids. The absence of patches of tomentum
on the inside of the tibiae would seem to exclude it from the closely
related Cylloepus.

Types are located in the Paris Museum and in the Francis Hunt-
ington Snow entomological collection. University of Kansas.

Sanderson: Species of Stenelmis 709

Stenelmis nevermanni Hinton

1936. Stenelmis nevermanni Hntn. Trans. Royal Ent. Soc. London, 85, pp. 424-426.

ORIGINAL DESCRIPTION

"Male. Elongate, subparallel, moderately convex. Surface
throughout microscopically alutaceous; above with the scutellum
glabrous, elsewhere sparsely clothed with very fine, moderately long
recumbent, pale-testaceous hair; elytra near apex with a few fine,
erect hairs which are often about as long as the combined length of
the two basal antennal segments; upper surface in many specimens
brownish-cinereous, due to a fine deposit of earthy material; beneath
clothed throughout with minute, dense tomentum and also clothed
as above. Derm pale brownish-piceous, feebly shining. Head with
the clypeal suture nearly straight; anterior margin of clypeus
truncate, with the angle on each side obtusely rounded; antennae
long and slender, extending slightly beyond basal prothoracic angles.
Surface very finely granulate, with the granules apparently seldom
separated by more than five times their diameters. Prothorax
longer than broad (30:26-5), broadest across basal one third; base
broader than apex (25:19). Apical margin feebly arcuate, feebly
sinuate on each side behind eye; apical angles inconspicuous, when
viewed laterally feebly produced forwards, obtusely rounded; sides
moderately converging towards apex, feebly, broadly sinuate in
apical one half and feebly, more narrowly sinuate before basal
angles, lateral margins indistinct, nearly obsolete, finely crenate due
to fine lateral granules; basal margin moderately strongly trisinuate,
broadly so on each side and narrowly so in front of scutellum.
Pronotum unevenly convex, without definite sublateral carinae;
median longitudinal impression feeble, extending from basal one
fourth to apical one third, deepest at apical one third. Surface
slightly coarsely, densely often confluently rugosely punctate; with
fine granules which are especially anteriorly and at sides separated
mostly by two to seven or more times their diameters. Elytra more
than twice as long as prothorax (68:30), broader across apical one
third than at humeri (39:35). Intervals nearly flat, alternate discal
intervals (1, 3, 5, and 7) slightly paler in colour than others; strial
punctures moderately coarse, subquadrate to round and on disk sep-
arated mostly by one to five times their diameters. Apices mod-
erately broadly but somewhat shortly produced, conjointly truncate.
Scutellum elongate, subovate, broadly rounded basally, acuminate
apically, length to breadth ratio is 6.5:3.8, flat, glabrous, shining.
Surface beneath finely granulate. Prosternal process as figured.

710 The University Science Bulletin

Metasterniim depressed posteriorly; with the median longitudinal
line extending only to anterior one third. Middle of first ventral
abdominal segment not noticeably depressed, granulate similarly to
metasternum, sides and other ventral segments much more finely
granulate than metasternum. Claws slender, not toothed. Length,
3.6 mm.; breadth, 1 mm.

"Female externally similar to male.

"Type: a male in my collection Costa Rica: Reventazon, 1. xi.
1934, at light (F. Nevermann).

"Paratypes: Twelve collected by the same collector at the same
locality during the following different times: — . xi. 1931, — , ii. 1932,
— . vii. and — . xi. 1934.

"Certain specimens appear to have a feebly developed fringe of
tomentum on the inner apices of the tibiae, but in all other respects
the species is a true Stenehnis. It is the only species of this cos-
mopolitan genus so far recorded from the Mexican or Central Ameri-
can region. It is close to none of the North American (U.S.A.)
species known to me."

Notes

Two specimens of this species from the type series have been
studied. Again this species is not closely related to those within the
United States, although it possesses certain characters which would
more closely associate it with the Crenata group. The last tarsal
segment is a little smaller than the preceding four combined, and
the claw is small and slender. The antenna is longer than the pro-
notum by the length of the last three segments. The third elytral
interval is but slightly elevated behind the extreme base, and the
sixth interval is not at all cariniform, as in the United States species.
The pronotum and venter are, in part, granulate, but the legs are
minutely punctulate. The extreme apices of the elytra meet at the
suture, and are squarely truncate.

This species, although agreeing with S. geayi Grouv. by its longer
antennae, is perhaps more closely related to the United States species.
It is smaller than S. geayi, slender, and is not variegated with
color.

The type and paratypes are located in the collection of Mr.
Howard E. Hinton, Berkeley, Cal. A paratype has been deposited
in the Francis Huntington Snow entomological collection. Uni-
versity of Kansas.

The two references below are to species which originally were
included in Stenelmis:

Sanderson : Species of Stenelmis 711

Cylloepus puncticollis (Hntn.)

]!)3t. Stenelmis puncticollis Hntn. Revista de Entoniologia, I\", pp. 198-1!»9.

I have not examined this species, but in a letter from its author,
it has been referred to Cylloepus. Cleaning and study of the an-
terior tibiae have shown the presence of tufts of tomentum on the
inner sides.

Microcylloepus pusillus (Lee.)

1852. Stenehnis pusillus Lee. Proc. Ac N. S. Phila. VI, p. 44.

1869. Limnius pusillus (Lee.) Zimmerman, Trans. Am. Ent. Soc, II, p. 259.

1870. Elmis pusillus (Lee.) Horn, Trans. Am. Ent. Soc. Ill, pp. 29-42.

1935. Microcylloepus pusillus (Lee.) Hinton, Stylops, Vol. 4, pt. 8, pp. 178-179.

The new genus Microcylloepus was erected by Hinton for this
and a number of related species.

Stenelmis canaliculata (Gyll.)

I have examined the type of Stenehnis elongata Mots.* which is
labeled: "Stenelmis elongatus Motsch. Am. bor.?" Careful com-
parison of the type with an authentically determined specimen of
the European Stenehnis canaliculata (Gyll.) convinces me that the
two are the same. S. elongata Mots, thereby becomes a synonym of
8. canaliculata (Gyll.) and should be dropped from our lists.

List of American Species of Stenelmis

NuBiFERA Group : page page

nubifera Fall 661 par\a n. sp 688

Crenata Group : fuscata Blatchley 689

sexlineata n. sp 663 hungerfordi n. sp 690

crenata (Say) 665 humerosa Mots 692

exigua n. sp 669 mirabilis n. sp 693

beameri n. sp 671 antennalis n. sp 695

lateralis n. sp 672 quadrimaculata Horn 696

concinna n. sp 674 musgravei n. sp 698

tarsalis n. sp 675 sinuata Lee 699

knobeli n. sp 677 decorata n. sp 701

bicarinata Lee 679 vittipennis Zimm 702

exilis n. sp 680 convexula n. sp 704

mera n. sp 682 miirkelii Mots 705

SINUATA-HUMEROSA G.iOUP : ^''^^'' ^'°''^' '^^

douglasensis n. sp 685 nevermanni Hntn 709

grossa n. sp 686

* Etud. Entom. VIII, 1859, p. 51.

712 The University Science Bulletin

LITERATURE CITED

Additional references, including those to fossil Dryopidae, will be found in
the text of this paper. (See table of contents.)

Barthe, E. 1927. Tableaux Analytiques des Coleopteres de la Faune Franco-
Rhenane (Dryopidae). Miscell. Entomologica, XXX, pp. 3-74.

Bertr.\nd, Henri. 1935. Larves de Coleopteres aquatiques de L'Expedition
Limnologique Allemande en Insulinde. Archiv f. Hydrobiologie. Suppl.
Bd. XIV, pp. 193-285, pi. II.

Bl.'^ckwelder, R. E. 1930. The Larva of Eubrianax edwardsi Lee. (Coleop-
tera, Psephenidae). Pan Pacific Ent.. \T, pp. 139-142, ill.

Blatchley, W. S. 1910. Catalogvie of the Coleoptera or Beetles known to
occur in Indiana, Indianapolis, pp. 1386, ill.

BoviNG, Adam G. 1929. On the Classification of Beetles According to Larval

Characters. Bull. Brook. Ent. Soc. XXIV, pp. 55-97.
BoviNG, A. G., and Craighead, F. C. 1930. An Illustrated Synopsis of the

Principal Larval Forms of the Order Coleoptera. Ent. Americana, XI

(n. s.), pp. 1-351.
Bradley, J. G. 1930. A Manual of the Genera of Beetles of America, North

of Mexico. Ithaca, N. Y., pp. 360.

Brocher, Frank. 1912. Recherches sur la Respiration des Insectes Aquati-
ques Adultes. Les Elmides. Annales de Biologie Lacustre (Tome V),
pp. 1-44, ill.

1913. L'Aquarium de Chambre. Paris, pp. 1-451, figs. 1-186.

Carter, H. J., and Zeck, E. R. 1929. A Monograph of Australian Dryopidae.
Australian Zoologist, Vol. VI, Pt. 1, pp. 50-72, PI. 7.

Darlington, P. J. 1936. A list of the West Indian Dryopidae (Col.) with a
new genus and eight new species, including one from Colombia. Psyche,
Vol. XLIII, Nos. 2 and 3, pp. 65-82, ill.

Dufour, Leon. 1835. Recherches Anatomiques et Considerations sur les
Insectes Coleopteres des Genres Macronique et Elmis. Ann. d'Sci. Nat.
Pt. Zo51., (2) 3: pp. 151-174, pis. 6 and 7, figs. 1-28.

Forbes, Wm. T. M. 1926. The Wing Folding Patterns of the Coleoptera.

Jr. of the New York Ent. Soc. XXXIV, pp. 42-138, ill.
Ganglbauer, L. 1904. Die Kafer von Mitteleuropa. Verter Band, erate

Halfte. 800. pp. 286. Wlen, Earl Ceroids Sohn.
HiNTON, H. E. 1935. Notes on the Dryopoidea, Col. Stylops, Vol. IV, Pt.

8, pp. 169-179.
• 1936. Notes on the Biology of Dryops luridus Erichson (Coleoptera,

Drvopidae). Trans, of the Society for British Entomologv, Vol. 3, pp.

67-78.

1936. Results of the Oxford University Expedition to Borneo, Dry-

opidae (Col.). Part I. Annals and Mag. of Nat. Hi.st. Ser. 10, Vol. XVIII,

pp. 89-109. Part II, 204-224.
Horn, G. H. 1870. Svnopsis of the Parnidae of the United States. Trans.

Am. Ent. Soc. Ill, pp. 29-42.
Lameere, a. 1900. Notes pour la classification des Coleopteres. Ann. Soc.

ent. Belg., 44:355-376.
LeConte, J. L. 1852. Synopsis of the Parnidae of the United States. Proc.

Ac. Nat. Sci. Phil., 6:41-44.
1861. Classification Col. N. Amer. pp. 114-117. Smithsonian Miscell.

Coll. III.
LeConte, J. L., and Horn, G. H. 1883. Classification of the Coleoptera of

North America, p. 568. Smithsonian Miscell. Coll. 507.

Sanderson: Species of Stenelmis 713

Leng, C. W. 1920. Catalogue of the Coleoptera of America North of Mexico,
J. D. Sherman, N. Y., pp. 470.

Matheson, Robert. 1914. Life History Notes on two Coleoptera (Parnidae)
Can. Ent. 46, pp. 185-189, ill.

SussKiND, M.ARiE E. CiMiNi. 1936. A Morphological Study of the Respiratory
System in Various Larval Instars of Stenelmis sulcatus Blatchley (Drj'-
opidae; Coleoptera). Papers of the Michigan Academy of Science, Arts and
Letters, Vol. XXI, published in 1936. Plates.

West, Luther S. 1929. A Preliminary Study of Larval Structure in the
Dryopidae. Annals Ent. Soc. of Amer. XXII, pp. 691-726, plates.

Zaitzev, p. 1908. Catalogue des Coleopteres aquatiques des families des
Dryopidae, Georyssidae, Cyathoceridae, Heteroceridae et Hydrophilidae.
Horae Societas entomologicae Rossicae. 37: pp. 283-420.

1910. Diyopidae, Cyathoceridae, Georyssidae, Heteroceridae. Coleop-

terorum catalogus auspicis et auxilio W. Junk. Pars 17.

Additional References Not Cited

The following references will be found useful to those engaged in a study
of Dryopidae :

Forbes, Wm. T. M. 1922. The Wing-venation of the Coleoptera. Annals Ent.
Soc. Amer., 15, pp. 328-352, 7 plates.

Leng, C. W. 1913. Aquatic Coleoptera, Jr. N. Y. Ent. Soc. XXI, pp. 32-42.
Mu.sgr-^ve, Paul N. 1935. Notes on Collecting Dryopidae (Coleoptera) Can.
Ent. LXVII, No. 3, pp. 61-63.

MuTTKowsKi, R. A. 1920. The Respiration of Aquatic Insects. Bull.
Brook. Ent. Soc. XV, pp. 89-96, 131-141.

Segal, Bernard. 1933. The Hind Wings of Some Dryopidae in Relation to
Habitat (Coleop.). Ent. News, 44: p. 85.

West, L. S. 1929. A Bibliography of the Drvopoidea. Suppl. to Battle
Creek College Bull. Ill, No. 1, pp. 3-12. (Contains 260 references.)

714 The University Science Bulletin

PLATE LXXX

Figs. 1 to 19. Male genitalia of Stenelmis.

Sanderson : Species of Stenelmis

715

2. S SEXUHCATA

PLATE LXXX

\J

6 S LATEPAUS

TSCONCINN*

6 S TAtiSAUS

10 S.BICAPINATA

16 S Mt/NGEftFO«OI

12 S OOUGLASCNStS

13 S (

15 S fuse ATA

i9.S.AflTLNNAUS

716 The University Science Bulletin

PLATE LXXXI

Fig. 1. Stenelrnis lateralis Sand. n. sp.

Figs. 2 to 5. Male genitalia of Stenelrnis species.

Figs. 6 and 7. Antennae of Stenelrnis species.

Fig. 8. Male genitalia.

Fig. 9. Middle tibia of female.

Fig. 10. Middle tibia of male.

Fig. 11. Male genitalia.

Figs. 12 and 13. Figures showing lower margin of last tarsal segment with
and without tooth-like projection.

Fig. 14. Tarsus of a species of Sinuata-humerosa group. The terminal
segment is equal to the four preceding combined; the claws are more robust.

Fig. 15. Tarsus of a species of Crenata group. The terminal segment is
shorter than the four preceding combined; the claws are more slender.

Fig. 16. Last visible ventral abdominal segment showing apical emargina-
tion.

Fig. 17. Figure showing well-developed hind wing.

Figs. 18 to 23. Outlines of types of elytral patterns in Stenelrnis species

Sanderson: Species of Stenelmis

717

PLATE LXXXI

2\.S.LAT£RALIS

23S

SexuntATA

D

17-4141

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