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Volume 1, Number 1

June 1, 1953

ON A NEW GENUS AND SPECIES OF MYSID FROM
SOUTHERN LOUISIANA

(Crustacea, Malacostraca)

ALBERT H. BANNER

DEPARTMENT OF ZOOLOGY AND ENTOMOLOGY,
UNIVERSITY OF HAWAII, HONOLULU

Ml GS&P. ZIBL
LI3KARY

IUL 1 4 1953

HARVARD
UNIVERSITY

TULANE UNIVERSITY
NEW ORLEANS

TULANE STUDIES IN ZOOLOGY is devoted primarily to the
zoology of the area bordering the Gulf of Mexico. Each number
is issued separately and deals with an individual study. As volumes
are completed, title pages and tables of contents are distributed to
institutions exchanging the entire series.

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Fenner A. Chace, Jr., Curator of Marine Invertebrates, United
States National Museum, Washington, D. C.

Mrs. Walter M. Tattersall, Pendeen, Sinah Lane, Hayling

Island, Hants., England.
George Henry Penn, Associate Professor of Zoology, Tulane

University of Louisiana, New Orleans, U. S. A.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
torial matters, and orders for individual numbers to the editor. Re-
mittances should be made payable to Tulane University.

Price for this number: $0.25.

George Henry Penn, Editor,

c/o Department of Zoology,
Tulane University of Louisiana,
New Orleans, U. S. A.

ms. comp.

fl 1 4 1953
HARVARD
UNIVERSITY

(Crustacea, Malacostraca)

ALBERT H. BANNER,

Department of Zoology and Entomology, University of Hawaii,

Honolulu.

Only five species of mysids have been reported from the fresh waters
of the Americas (Tattersall, 1951:4; Banner, 1948:72). Three of
these are known only from tropical America: Antromysis cenotensis
Creaser, from the caves of Yucatan; As anophelinae Tattersall, from the
holes of a terrestrial crab in Costa Rica; and Diamysis americana
Tattersall, from ditches in Dutch Guiana. The other two species are
found in temperate and arctic North America: My sis relic ta Loven,
which reaches south from the arctic coast to the Great Lakes, and
which apparently is a circumarctic species; and Neomysis mercedis
Holmes,2 which ranges from brackish into fresh waters along the
Pacific Coast from California to Alaska. No mysids have been re-
ported before from the fresh water of the Mississippi basin.

TAPHROMYSIS 3, gen. nov.

Diagnosis — Body of usual form, carapace covering most of thorax.
Eyes well developed. Antennules of normal configuration. Antennal
scale rounded and bearing setae on both sides. Labrum rounded an-
teriorly. Mandibles with strong dentate process laterally on corpus
mandibulae. Tarsus of thoracic legs (e.g., portion beyond "knee") of
four articles. Three pairs of oostegites in females, but those of the
sixth thoracic segment strongly reduced. Adult males without sternal
thoracic processes. All female pleopods reduced to plates; first and
second pleopods of males similar. Third pleopods of males consisting
of two articles, representing the protopod and endopod; exopod lack-
ing. Fourth pleopods of males large, with exopod of seven articles,
and bearing strong terminal bristles that form a feeble "pincher";
endopod short and composed of two articles. Fifth pleopod of males
of two slender articles. Uropods without spines near statocyst. Telson
posteriorly cleft.

Type species: Taphromysis louisianae Banner.

Discussion — Taphromysis was erected solely for the species de-
scribed below.

This genus plainly belongs to the family Mysidae, the sub-family
Mysinae and the tribe Mysini of H. J. Hansen (1910:13), and within

1 Contribution No. 35, Hawaii Marine Laboratory.

2 The taxonomy of this species will be discussed in another
paper.

3 Generic name derived from taphros, Greek for ditch, refer-
ring to the habitat where the type species was found.

4 Tulane Studies in Zoology Vol. 1

the tribe to the Mysis-group of Zimmer (1915:202-216). This group
may be recognized by a series of characteristics — the lack of an an-
terior process on the labrum, the possession by the males of six or
seven articles in the exopod of the fourth pleopods, and the develop-
ment on this ramus of long terminal and sub-terminal bristles to form
a type of a feeble "pinchers"; and the possession of a truncate or
terminally cleft telson. This group until now contained six recognized
genera: My sis Latreille, Hemimysis G. O. Sars, Par amy sis Czerniavsky,
Caspiomysis G. O. Sars, Schistomysis Norman and Synmysis Czerniav-
sky. From the last four genera Taphromysis is separated by a number
of characteristics, the most important being the absence of the exopod
on the third pleopod of the males and the presence of setae on the
outer margin of the antennal scale.

Taphromysis shows many similarities to Mysis and especially to
the species M. relicta Loven. If female specimens alone were avail-
able to study, only the shorter antennal scale and the presence of the
mandibular process could be relied upon to distinguish Taphromysis.
In the male, however, the exopod of the third pleopod is well de-
veloped and consists of six articles in Mysis while the entire ramus
is lacking in this genus. A difference of this category in the present
scheme of classification is considered to be of generic worth.

Taphromysis also shows close affinity to the genus Hemimysis,
The antennal scale, variable in Hemimysis, usually bears setae only
on the distal portion of the outer margin instead of along its entire
length as in this genus. The presence of the dentate process on the
lateral surface of the mandible has not been noted in Hemimysis.
The oostegites are of similar development in the two genera. In
Hemimysis the exopod of the third pleopod of the males may be
reduced or entirely wanting as it is in this genus. The fourth pleo-
pods of the males are the same in both genera [according to Illig
(1950:587), but Tattersall and Tattersall (1951:331) state that the
exopod in Hemimysis consists of only five to six articles] . The telson
in both genera is quite similar. However, the two genera may be
distinguished by the fifth pleopods of the males which in Hemimysis
are well-developed, biramous and natatory, but are reduced to two
simple elongate articles in Taphromysis.

While this genus appears to lie midway between Mysis, to which
it is similar in the nature of the fourth and fifth pleopods of the
male, and Hemimysis, to which it is similar in the nature of the third
and fourth pleopods of the male, the modification of the third pleo-
pods appears to be more distinctive than that of the fifth, and the
genus, therefore, probably is more closely related to Hemimysis.
However, it appears to me that these specimens raise doubts as to
the validity of small modifications of the antennal scale and of the
pleopods of the male as characteristics for generic separations.

It is noteworthy that the American Taphromysis should be so
closely related to the genus Hemimysis which is restricted to the
waters of western Europe and the general Mediterranean region,

No. 1

Banner: New Mysid from Louisiana

penetrating into the brackish water of the Black Sea drainage of
Rumania (Bacesco, 1940). Tattersall (1951:228) has postulated
that the genus Diamysis, a brackish water genus with three Mediter-
ranean species — ecologically similar, therefore, to Hemimysis — and
one American fresh water species, was once a widespread tropical
genus of the Tethys Sea of the Tertiary, and that the modern discon-
tinuous distribution represents a relict fauna. It is possible that the
genus Hemimysis once had a similar distribution, but in the case of
its American species the long isolation resulted in sufficient modifi-
cation to support its classification as a separate genus.

10 mm

Figure 1. Taphromysis louisianae, adult male.

TAPHROMYSIS LOUISIANAE, sp. nov.

Type specimens — A male 7.0 mm long, United States National
Museum, Catalog No. 94829; paratypic series at the United States
National Museum, Catalog No. 94830 and also in Tulane University
collection, No. 2643.

Type and paratypes all collected in a roadside ditch at Gueydan,
Vermilion Parish, Louisiana, Feb. 2, 1952 by F. F. Vizzi. The water
was reported to be fresh and static; the bottom was reported to be
mud and the vegetation to be sparse.

6 Tulane Studies in Zoology Vol. 1

Description — Adult females about 8.0 mm long, adult males up to
7.5 mm long.

Body of usual development and proportions. Carapace posteriorly
leaving dorsal portion of the eight and part of seventh thoracic seg-
ments exposed. Rostral plate short, scarcely reaching the bases of
eyes, rounded and depressed. Anterior margin of carapace slightly
below base of eyes bearing small acute tooth. Anterior lateral corners
of the carapace rounded.

Eyes somewhat elongate and sub-cylindrical, with total length twice
maximum breadth of stalk and with evenly rounded corneas 0.4 length
of entire stalk. No ocular papilla.

Antennular peduncle with end of second article reaching slightly
beyond end of eyes. Basal article 2.7 times length of second and 2.0
times length of third articles. No article bearing any conspicuous
teeth or denticles. Process masculinus well developed, 1.4 to 2.0
times length of third antennular article, and bearing fine setae. Outer
flagellum slightly longer than body, inner flagellum about half body
length.

Basicerite of antenna bearing small lateral tooth. Scale reaching
to end of antennular peduncle, about 4.6 times as long as broad,
bearing setae on both sides; tip rounded; distal twentieth demarked
by feeble articulation. Carpocerite of antennal peduncle reaching past
middle of scale. Flagellum as long as outer antennular flagellum.

Labrum of normal form, anteriorly rounded.

Opposing faces of mandibles with usual development (rows of
spines imperfectly shown in drawings because of the rotation of
appendage). Corpus mandibulae with strong acute denticle laterally
directed. Palp normal, maxillules and maxillae normal.

First thoracic leg (maxilliped) with exopod and endopod located
relatively close together on the protopod, but not adjacent. Endite
of basis and two basal articles of endopod of moderate development.
Exopod only slightly shorter than that of second leg. Second thoracic
leg also of normal development.

Third to eighth thoracic legs with basis somewhat enlarged; tarsus
(or portion beyond "knee") of four articles, including the "end-claw";
penultimate article bearing heavy spine, more than twice as long as
end-claw. All exopods well developed with outer distal angle of
basal plate sub-acute and not bearing accessory denticles.

Penis lobes of males well developed. Females bearing three pairs
of oostegites, with those of sixth thoracic legs reduced to small seti-
ferous lobes.

All pleopods of females reduced to uniramous plates. First and
second pleopods of males similar to female pleopods. Third male
pleopod with enlarged protopod, exopod entirely lacking, endopod
longer than protopod but less than half as broad, consisting of a
single article. Protopod and endopod bearing setae.

No. 1

Banner: New Mysid from Louisiana

Figure 2. Taphromysis louisianae, adult male, a, anterior body region, dorsal
aspect; b, antennular peduncle, medial aspect; c, antennal peduncle and
scale, ventral aspect; d, mandible, posteromedial aspect; e, mandible,
medial aspect; f, maxilla; g, first thoracic leg; h, second thoracic leg;
i, seventh thoracic leg; j, third pleopod ; k, fourth pleopod ; 1, tips of
distal setae of fourth pleopod; m, fifth pleopod; n, uropod ; o, telson.
(Figures d and e, scale B; figure f, scale C; figure 1, scale D; all other
figures scale A.)

8 Tulane Studies in Zoology Vol. 1

Fourth pleopod of males very elongate, reaching to distal tip of
uropod. Basal article of protopod with heavy but short lobe on
outer side. Second article reaching beyond end of fifth abdominal
segment. Endopod consisting of two articles, basal only one seventh
as long as distal and bearing small exite; distal article reaching beyond
second article of exopod and bearing numerous setae. Exopod com-
posed of seven articles, with the articulations between them very faint.
Penultimate article bearing a curved spine, as long (if straightened)
as the exopod to its base; bearing distally a series of spiral reinforce-
ments and a series of extremely fine spinules on its acute tip.
Terminal article fused with strong terminal seta, more blunt and
straight but slightly shorter than penultimate setae, distally armed
with very fine spinules in diagonal bands. Terminal and subterminal
setae forming a feeble "pincher."

Fifth pleopod of male with tip reaching almost to end of sixth
abdominal segment, composed of two articles, the distal about 1.5
times length of proximal. Setae found only on distal portion of
second article; terminal setae as long as appendage.

Telson short, less than 0.9 as long as sixth segment, broad and
distally emarginate. Tip about 0.6 as broad as base, and 0.4 as broad
as telson is long. Lateral margins concave and bearing 10 pairs of
movable spines. Posterolateral spines not longer than lateral spines.
Terminal emargination less than 0.2 length of telson, rounded and
"U" shaped, and bearing about twenty pairs of long narrow denticles
that are not articulated.

Uropods slightly less than twice as long as telson, of usual form
and not bearing any spines near statocyst.

Specimens in preservative without color but bearing a limited
number of small stellate melanophores, one middorsally on each ab-
dominal segment, a pair on base of telson, several pairs on ventral side
of cephalothorax and its appendages, and two pairs on oostegites.

REFERENCES CITED

Bacesco, M. 1940. Les Mvsidaceas des eaux Roumaines . . . Univ.
de J assy., Ann. Sci., 2nd Ser., 24(2): 453-804.

Banner, A. H. 1948. A taxonomic study of the Mvsidacea and
Euphausiacea (Crustacea) of the Northeastern Pacific. Part
II. . . . R. Canad. Inst, Trans., 27(57): 65-125.

Hansen, H. J. 1910. The Schizopoda of the Siboga Expedition.
Siboga-Exped., 37: 1-123.

Illig, G. 1930. Die Schizopoden der Deutschen Tiefsee-Expedi-
tion. Wiss. Ergebn. Deutschen Tiefsee-Exped., 22(6): 399-625.

Tattersall, W. M. 1951. A review of the Mysidacea of the
United States National Museum. U. S. Natl. Mus., Bui., 201:
x +292.

Tattersall, W. M. and O. S. Tattersall. 1951. British Mysi-
dacea, viii +460. Ray Society, London.

Zimmer, C. 1915. Die Systematik der Tribus Mysini H. J. Han-
sen. Zool. Anz. 46: 202-216.

Volume 1, Number 2

July 3, 1953

A CONTRIBUTION ON THE LIFE HISTORY OF THE LIZARD
SCINCELLA LATER ALE (SAY)

RICHARD M. JOHNSON

DEPARTMENT OF BIOLOGY, TENNESSEE WESLEYAN COLLEGE,

ATHENS, TENN.

MUS. C08P. 208L
LSB3ARY

AUG IMI58

UNIVERSITY

TULANE UNIVERSITY
NEW ORLEANS

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Hobart M. Smith, Associate Professor of Zoology, University
of Illinois, Urbana.

Henry S. Fitch, Associate Professor of Zoology, University of
Kansas, Lawrence.

Fred R. Cagle, Professor of Zoology, Tulane University of

Louisiana, New Orleans.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.
Address all communications concerning exchanges, manuscripts, edi-
torial matters, and orders for individual numbers to the editor. Re-
mittances should be made payable to Tulane University.

Price for this number: $0.75.

George Henry Penn, Editor,
c/o Department of Zoology,
Tulane University of Louisiana,
New Orleans, U. S. A.

A CONTRIBUTION ON THE LIFE HISTORY OF THE LIZARD
SCINCELLA LATER ALE (SAY) *

RICHARD M. JOHNSON 2

Department of Biology, Tennessee Wesleyan College, Athens, Tenn.

The objectives of this research were threefold: to determine
size at sexual maturity, the seasonal reproductive cycle, and the repro-
ductive potential of the scincid lizard Scincella later ale (Say). Data
to accomplish these objectives were obtained by laboratory examination
of fresh and preserved material, the latter from the research col-
lections of the Department of Zoology of Tulane University.

Blount (1929), Breckenridge (1943), and Reynolds (1943) pre-
sent data obtained from histological examinations of gonads of Phry-
nosoma solare, Eumeces s. septentrionalis and Eumeces jasciatus re-
spectively. Each of these lizards exhibits essentially the same morph-
ological and physiological sequences of the gonads. The appearance
of spermatozoa in January and February is associated with growth of
the testes to maximal size; spermatozoa are present in the gonadal
ducts through August; disappearance of spermatozoa is concurrent
with regression of testes to minimal size in October and November.
The ovaries follow the same sequences except for a more rapid en-
largement in early spring and a more sudden regression in late sum-
mer. Scincella laterale parallels the above lizards as regards these
phenomena.

MATERIALS AND METHODS

Four hundred and sixty-two specimens (229 males, 233 females)
were examined during this investigation. The majority of the speci-
mens had been preserved prior to this study.

Klauber (1943: 20) discusses shrinkage of linear dimensions of
snakes in preservative. As a standard procedure is observed for en-
tering specimens in the Tulane University collections, it is assumed
that the amount of shrinkage for the same parts of different in-
dividuals was the same, if any. If such an assumption is true, then
experimental error would not be significant (Simpson and Roe,
1939).

Data obtained from each lizard were: measurements of axilla-groin
and snout-vent lengths to the nearest 0.1 mm; counts of dorsal and
midbody scale rows as prescribed by Smith (1946: 27, 29, 30);
counts and measurements of ovarian follicles and oviducal eggs; the
absence or presence of spermatozoa in the testes and/or epididymis
and vas deferens.

1 The generic name Scincella follows the nomenclature of Mittie-
man (1950).

2 A thesis submitted in p^WflSl f&^^hMJjf""^ the requirements
for the degree of Master of Science '©^ffie,\pfritHifte School of Tulane
University.

AUG 2 4 io^j
HARVARD

12 Tulane Studies in Zoology Vol. 1

Testes and gonadal ducts were crushed on a slide and examined
under a compound microscope. Filamentous structures converging
from the periphery to the center of seminiferous tubules were in-
terpreted as tails of spermatozoa. Tadpole-like structures in the
epididymis and vas deferens were accepted as evidence of mature
spermatozoa. The presence of spermatozoa in either the testes or
gonadal ducts was accepted as a criterion of sexual maturity (Cagle,
1948a: 108; 1948b: 1). Sexually active males are those containing
spermatozoa in the epididymis and vas deferens (Cagle, 1948b: 1).

Measurements of ovarian follicles and oviducal eggs were obtained
with an ocular micrometer. The former were measured for greatest
diameter, the latter for greatest length and width. After being mea-
sured each egg was opened for inspection. All oviducal eggs observed
during this study were enclosed in leathery shells with fine longi-
tudinally oriented striations.

Tail lengths were used in the study of sexual dimorphism of males
and females of series Tulane 14108. Only tails showing no interrup-
tion of symmetry of scalation, except for deletion of scale rows, and
not showing signs of regeneration were considered. Because of the
high frequency of incomplete or regenerated tails in series Tulane
14108, this measurement was subsequently abandoned.

Of the 32 males and 37 females in this series, 20 males (62.5%)
and 23 females (62.2%) had broken or regenerated tails. There
was no indication of correlation between snout-vent length and the
frequency of tail injury. The loss of the tail may be ascribed to injury
as a result of contact with members of the same species (Carr, 1940;
Lewis, 1951) and/or predation. This author has observed captives
of Scincella seize and attempt to eat the tails of cage mates.

To obtain estimates of growth rates, size groups of 4 mm were
selected. The range between minimum and maximum snout-vent
lengths of hatchlings was 4 mm. The bias of collecting and possibly
the method of selecting size classes may introduce error in the esti-
mate of growth rate.

The method of plotting Figures 1-5, 7-10, is adapted from Cazier
and Bacon (1949). In these figures the horizontal bar is the mean,
the solid rectangle is the mean plus-and-minus three standard errors, the
dashed line the mean plus-and-minus three standard deviations, and
the solid line the range of observations. Abbreviations and statistical
symbols employed are: b, regression coefficient; m, sample mean; n,
number of observations; s, standard deviation; sm, standard error
(Snedecor, 1946); T, value of significance of difference between
any two statistics (Peatman, 1947).

SEXUAL DIMORPHISM

Series Tulane 14108 from Bonnet Carre Spillway, St. Charles Par-
ish, Louisiana, collected February 12, 1950, was used as the standard
to determine sexual dimorphism. The following data were employed:

No. 2

]ohnson: Life History of Scincella

snout-vent lengths; axilla-groin/snout - vent length; snout - vent/tail

Hales Females

Hales

Females

T
. i _

1
1

T ' T '

. 1
1

-L

30 .

*£

J. 1

i 1

n 31

29 36

m 67.5

73.0

30.1 30.0

fm -51

.22 .18

s 2.86

3.23

1.20 1.07

T 8.1

0.00

Compart

of Dors*

Tulane

rig. l

ion of the Bunker
il Scale Howe of
Series 1*»10B.

Fig. 2.

Comparison of the Number

of Mldbody Scale Howe of

Series Tulane 1>*108

Hales

Females

.635

.605

.575

.51*5

.515

.1*91

.513

•.003

.017

6.2

Hales

Females

.57k

.ool*

.022

is. 3.

Comparison of Axilla-
Oroln/Snout-Vent Hatlos
of Series Tulace 11*108.

.730

.670

.610

.550

.1*90

.570
.005
.019
3.5

Fig. K

Comparison of Snout-
Vent/Tall Length Ratio*
of Series Tulane 1U108.

Kales

Females

+ t

I
I
-L

1*2.7

.63

3.58

1.3

1*3.8
.67
lt.06

n«. 5.

Comparison of Snout-
Vent lengths of Serlee
Tulane 1>*108

14 Tulane Studies in Zoology Vol. 1

length; dorsal and midbody scale row counts. Pigmentation of the
gular region and lower labials was also studied.

These males and females differ significantly [T greater than 2.5
(Peatman, 1947)] with respect to dorsal scale row counts, axilla-
groin/snout- vent length ratios, and snout- vent/tail length ratios (figs.
1, 3, 4). The larger values for dorsal scale row counts and axilla-groin/
snout-vent length ratios for females might be correlated with capacity
for egg production as suggested for snakes by Blanchard (1921: 6)
and Ortenburger (1928: 10-11). The males and females did not dif-
fer significantly with respect to midbody scale row counts and snout-
vent lengths (figs. 2, 5), but the females possessed the greatest snout-
vent length. All of these specimens are 35 mm or more in snout- vent
length. It is evident that sexual dimorphism does exist for specimens
of this size with males having fewer dorsal scale rows, smaller axilla-
groin/snout-vent ratio, and smaller snout- vent/ tail length ratio (i. e.,
longer tails).

Identification of sex, verified by dissection, is not possible on the
basis of pigmentation. Gular pigmentation is most intense in those
individuals having the darkest overall coloration.

Klauber (1937: 12, 16; 1943: 31) has shown sexual dimorphism
to exist in juveniles of certain species of snakes. To test for sexual
dimorphism in juvenile Scincella, a series of 16 males and 8 females
less than 35 mm snout- vent length were utilized. As discussed below,
35 mm snout-vent length is the lower limit of sexually mature individ-
uals. The males ranged from 16.5 mm to 34.8 mm, the females from
18.9 mm to 34.8 mm. For the character of axilla-groin/snout-vent
length ratio these males and females could be considered significantly
different, T=2.4; males: m, .489, sm, .003; females: m, .526, sm, .003.
The T-value is probably high as a result of the small numbers of
individuals compared and of the difference in average age as indicated
by the difference in average snout-vent lengths; males: m, 22.4
mm, sm, .45 mm; females; m. 27.2, sm, .66 mm; T=2. Sexual di-
morphism is shown in figure 6 to begin at about 24 mm snout-vent, but
is not definitely shown by the series under discussion. Rather, this
series may show changed body proportions in young of different age
groups. The inclusion of nearly mature individuals in the upper limits
of snout- vent length may also contribute to this apparent difference
(Klauber, 1943). These males and females differed significantly as
regards dorsal scale counts: male, m, 68; females: m, 74, T=3.3, but not
from the counts of males and females respectively of series Tulane
14108. They did not differ significantly from each other, nor from
males and females of the latter series, as regards midbody scale row
counts.

Coefficients of regression of axilla-groin on snout-vent of 66 males
and 85 females are .93 and .98 respectively (fig. 6). Lewis (1951)
and Smith (1946) do not give criteria for distinguishing sexes.

Descriptions of Scincella refer to it as a smooth-scaled lizard. Scales

No. 2

Johnson: Life History of Scincella

on the sides of the neck, ventrolateral body surfaces, posterior limb
surfaces, and the basal tail region each possess three to four small
keels. A few specimens also have keels on the dorsal scales. No keels

Axilla-
groin

Males, 1- -2.7 .6U

Snout-vent

Males ° Females

Axilla-groin Snout-vent Axilla-groin Snout-vent
n 66 85
m 19.8 37.1 22.8 1,0.1*
s^ .65 1.04 .72 1.04

a 5.29 8.43 6.60 9.58

Tig. 6.

Correlation of ixllla-Groia to Snout-Tent Lengths of Males and females

occur on the scales of the venter. The sexes cannot be distinguished
by the absence, presence, or distribution of these keels. The only
correlation suggested was with age. The smallest individuals possess
the most conspicuous keels and are most frequently the individuals
with keeled dorsal scales.

GEOGRAPHIC VARIATION

The wide distribution of Scincella offers the possibility for geo-
graphic subspeciation (Smith, op. cit.). Mittleman (1947) considers
the Mississippi River and Lake Pontchartrain in Louisiana as a line
of demarcation between subspecies of the salamander Manculus quad-
ridigitatus. Samples of Scincella from selected areas within the state
were compared for possible geographic variation in certain morpho-
logical characters.

Three groups of specimens were compared. These were selected for
homogeneity of size class, sex, and locality of collection. The 69 speci-
mens, series Tulane 14108, described above were used as the standard
to represent central Louisiana. A series of 25 males and 22 females
from Grand Isle, Jefferson Parish, and Plaquemines Parish, was se-
lected to represent southern Louisiana. Fourteen males and 11 females
from adjacent parishes north of Lake Pontchartrain were selected as
the northern sample. These series were compared, sexes separately,

Tulane Studies in Zoology

Vol. 1

with respect to axilla-groin/snout-vent length and dorsal scale row
counts (figs. 7-10) and midbody scale rows. Tests for significance of
difference were negative for these characters among the three regions.
Midbody scale row counts, means plus-and-minus three standard errors,
of the northern and southern samples, [males, 29.1 ±.32 and 29.4±.27
respectively, and females, 29.2±.30 and 30.2±.37 respectively] are
well within the limits of the males and females of series Tulane 14108

Southern

Bonnet
Carra

.590

.570

.550

.530

.510

•U»o

Bonnet

Carre

Northern

i ( (

t i

9
Jo

)

lit

.551

.5U3

.551

.003

.001*

.016

.017

.015

Conparlcon of
Ratio.

PI
Ajllla-
jf Three

S. 7.

Grota/Saout
Serl«B of

-7«ot

Haiti

Uaitb

Southern

BoDnet
Carro

riorthern

1 X

T t' '

* * t

25
72.3
.52
2.61

73.0
.54
3.23

73.9
.76
2.54

Tig. 10.

Comparltoo of the Sumter of Doreal Scale Howe
of Three Serlee of Teoalee.

I
I
X

66.5

.72

3.36

67.5

.51

2.36

rig. 8.

Cooparlloa of the Buober of Doreal Scale Bowe
of Three Serlee of Hale*.

Southern

Bonnet
Carre

.6U0

.600

.560

.520
.508

northern

T
I

I 4

25
.577
•OOll
.022

J.
37

•57li
•OOU
.022

i I

.582
.005
.018

««. 9.

Cenparlsoo of ixllla-iroln/Snout-Veol Length
Batloe of Three Serlee of fenalee.

(fig. 2). Lewis, op. cit., lists 30 midbody scale rows and 61-71 dorsal
scale rows for Texas specimens. These counts are in agreement with
Louisiana specimens if the counts of the dorsal and midbody scale
rows of males and females are combined.

No. 2 Johnson: Life History of Scincella 17

Coloration was of no value in these comparisons as it is difficult to
evaluate and showed no subjective geographic variation. Specimens
from all three areas could be matched as regards extremes of intensity
and color pattern.

As regards the morphological characters studied, the Louisiana
Scincella may be considered as a single form. Thus, series of lizards
from various parts of the state may be combined for purposes of this
study. As there is a decided difference in the ecology of northern and
southern Louisiana (Viosca, 1933), the possibility of physiological dif-
ferences remains (Mayr, 1942).

SEXUAL MATURITY AND SEASONAL ACTIVITY

Males contained mature spermatozoa in the testes at a minimum
class range of 32-35 mm (actually 35 mm) snout-vent length (fig.
11). Spermatozoa are present in the testes and/or gonadal ducts from
January through August. Sexual activity thus prevails for eight months
of the year. The absence of spermatozoa from the ducts, but their
presence in the testes, of individuals in the lower class limits in Feb-
ruary and March indicates that these individuals are entering their
first season of sexually maturity. Complete absence of spermatozoa in
the sexually mature size group individuals during the breeding season
(January through August) occurred in only four instances. A February
specimen is probably immature. The other three, two in March in
the 40-47 mm size groups, one in August in the 44-47 mm size group,
may be senile or otherwise physiologically incapacitated. The August
specimen may have exhausted its spermatozoa. Spermatozoa are ab-
sent from all individuals of the sexually mature size groups from
October to December. There are too few specimens from the month
of September for any assumptions.

On the basis of the presence of oviducal eggs, the minimal size
group for sexually mature females is 40-43 mm (actually 40 mm).
Only two instances (December) of ovarian follicles equal to or greater
than 1 mm in diameter are present during the period of sexual inacti-
vity of males (October through December). On the basis of possess-
ing follicles of 1 mm or more in diameter during the breeding season
(cf. discussion of reproductive potential) females may be considered
mature. The minimal size group exhibiting this criterion is 36-39
mm snout-vent. No individuals less than 35 mm snout-vent possessed
follicles of this diameter at any time during the year. The presence of
follicles of this diameter coincides with the presence of spermatozoa
in the vas deferens of males. Oviducal eggs are present during the
months of March through August, indicating potential egg deposition
for those months. Literature reports of earliest dates for oviducal eggs
are March 25, Mississippi (Cook, 1943: 19) and April 7, Texas
(Lewis, op. cit.).

Assuming copulation to be concurrent with the presence of sperma-
tozoa in the vas deferens and of ovarian follicles of 1 mm or more in

Tulane Studies in Zoology

Vol. 1

April

2 3 6 A 2

1 1

3 9 2

October

May

A 3

November

3
June

13 9^

13 3 5 3 1

December

CM oj m c^n f\ ->t -^ vn

^O O -4 «0 CM -A O -* TO

v0O-V*0cl\0©.A«4

Snout-vent lengths (mm)
Spermatozoa absent 1 I Spermatozoa in testes r>\M Spermatozoa in ducts

Fig. 11.
Seasonal Distribution of Kales

January

1 1 A 1

1 2 2 A

July,

K 5 1

February

6 17 15 9

15 5 3 11

August

W///////A

gaga

March

12 5 9 3

September

1 2 A 5 5

L',

Hay

November

, , 1

15 8

2 1

1 2

June

rrrkv^V

December

1 6=5=51

ciii i j i i i i i i i i i

»■ »y ih *» »» *i

^o o -** co cv no

3 3 $

Snout-vent lengths (mm)
Follicles 1 mm. or more in diameter P^?l Oviducal eggs present KS3

Fie. 12.
Seasonal Distribution of Females

No. 2 Johnson: Life History of Scincella 19

diameter, a time lapse is indicated between copulation and ovulation
(figs. 11, 12). It is established that copulation and ovulation are not
concurrent in other reptiles (Blount, 1929; Breckenridge, 1943; Cies-
lak, 1949). Breckenridge, op, cit., reports 35 days as the time between
the last observed copulation and the first appearance of eggs. He does
not specify oviducal or deposited eggs.

Only one instance of possible copulation was observed in the field.
Copulation was not observed in the laboratory. On July 25, 1950, two
Scincella were discovered beneath a board in a position similar to that
pictured for E. s. septentrionalis (Breckenridge, op. cit., 596). The
position was retained for approximately 30 seconds after their dis-
covery before they separated and ran into the grass. Reports of copu-
lation were not found in the literature.

The earliest date for field collected eggs in Louisiana is May 12
(for this study), and is the earliest known date for any region. The
latest date for field collected eggs in Louisiana is July 23. The oc-
currence of oviducal eggs in August indicates possible deposition in
September in Louisiana. Cook, op. cit., and Lewis, op. cit., record
clutches for August in Mississippi and Texas respectively.

REPRODUCTIVE POTENTIAL

Determination of the number of broods per female per season re-
quires examination for oviducal eggs and of ovarian follicles. There
was a total of 31 females with oviducal eggs from March through
August. Two contained eggs only in the right oviduct. No female had
eggs only in the left oviduct. The range of numbers of eggs per fe-
male was from one to five (two eggs in the left oviduct, three eggs
in the right). The average number, plus-and-minus one standard
error, of eggs for the right and left oviducts respectively is 1.9±.06 and
1.5±.05. This difference is significant, T=5.1. The average num-
ber of eggs per female is 3-3— .05. The range of numbers of eggs
per clutch recorded in the literature is from one to five.

A significant correlation of number of eggs to snout-vent length per
female does not exist, £=.069 (fig. 13). Neither is there a correlation
between average egg length and snout- vent length (fig. 14).

During the months of October through December, only two
females contained ovarian follicles of a maximum diameter of 1
mm or more. Each of these individuals was collected in December. A
female 49-3 mm snout-vent had follicles 1.09 mm and 1.13 mm in dia-
meter; a female 38.8 mm snout- vent had a follicle 1.01 mm in dia-
meter. (The follicle diameters above and subsequently reported are
the maxima of left and right ovaries.) The maximum diameter of a
follicle of a sexually mature female for October was 0.92 mm. The
smallest follicle of maximum diameter of a female of sexually mature
snout- vent length was 0.57 mm (November). The smallest female,
27.2 mm snout-vent, collected in November had a maximum follicle
diameter of 0.50 mm. The average maxima, October, November, De-

Tulane Studies in Zoology

Vol. 1

cember, follicle diameters (right and left maxima combined) were:
0.79 mm, 0.80 mm, and 0.94 mm respectively for females 35 mm or
more snout-vent, and 0.59 mm, 0.66 mm, and 0.85 mm respectively

CO
bD
bf>
CD

b = .069

40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57
Snout-vent length (mm)

Correlation of Numbers of Ovlducal Eggs with Snout-Vent Lengths

bO
0)

«H

b = .009

40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57

Snout- vent length (mm)

Correlation of Average Lengths of Sggs with Snout-Vent Lengths.

for females less than 35 mm snout- vent. The greater monthly average
increase in follicle diameter for the latter group of females is probably
a reflection of the approach of larger individuals to sexual maturity.
Data are too few for September and January.

February collections contained no individuals with oviducal eggs, but
the largest follicle diameters were observed among these individuals.

No. 2 Johnson: Life History of Scincella 21

The largest follicles observed for February were in females 50.9 mm
and 51.4 mm snout- vent. Respectively, these follicles were: 3.90 mm,
4.23 mm, and 4.50 mm, 4.66 mm in diameter. Presumably these are
near ovulation. The smallest maximum follicles observed in sexually
mature females were: 0.74 mm, 0.88 mm, and 0.77 mm, 0.92 mm
respectively in females 38.9 mm and 39.8 mm snout-vent. The aver-
age maximum follicle diameter for February was 1.33 mm. No fe-
males less than 35 mm snout- vent were available for study.

March, April, and May are treated together because only one in-
dividual less than 35 mm snout-vent is present (April) and because
oviducal eggs were present. The largest follicles (March) were those
of two females 49.0 mm and 52.0 mm snout- vent. Respectively, these
follicles were: 4.77 mm, 4.43 mm and 3-20 mm, 3.47 mm. One fe-
male contained two eggs in each oviduct. This female, 49-9 mm snout-
vent, had the following follicle diameter maxima: 1.91 mm, 1.79 mm,
1.57 mm, and 3.70 mm, 3.05 mm. The smallest follicle diameter max-
ima for March were: 0.88 mm, 0.84 mm (female, 45.5 mm) and 0.80
mm (female, 49.8 mm). For April, the largest follicles were: 3.48
mm, 3-58 mm (female, 40.1 mm). The smallest follicle maxima for
April were: 0.78 mm, 0.80 mm (female, 34.8 mm). Two females
(47.8 mm and 50.6 mm snout-vent) contained oviducal eggs in
April. Respectively, their follicle maxima were: 1.01 mm, 1.05 mm
and 1.32 mm, 1.30 mm. Follicle diameter maxima for May were: 3.50
mm, 3-68 mm (female, 47.8 mm). Two females collected in May had
oviducal eggs and had the following follicle diameter maxima: 1.27
mm, 1.34 mm (female, 49.4 mm) and 1.09 mm, 1.16 mm (female,
46.6 mm). The smallest follicle maxima were those of the latter fe-
male above. The average follicle diameter maxima for March, April,
and May respectively, were: 1.60 mm, 1.68 mm, and 3-09 mm.

In the June, July, and August collections, four, four, and two fe-
males respectively contained oviducal eggs. These females had the
following follicle diameter maxima: 2.12 mm, 2.22 mm (female, 44.9
mm), 1.72 mm, 1.76 mm (female, 47.0 mm), 2.49 mm, 2.52 mm
(female, 44.3 mm), 1.76 mm, 1.91 mm (female, 46.3 mm) for
June; 1.26 mm, 1.42 mm (female, 45.2 mm), 2.59 mm, 2.66 mm
(female, 50.8 mm), 3.10 mm, 3.26 mm (female, 48.2 mm), 1.15 mm,
1.26 mm (female, 53.7 mm) for July; 2.20 mm, 2.29 mm (female,
56.8 mm), 1.11 mm, 1.15 mm (female, 52.2 mm) for August. The
largest follicle diameter maxima for females not containing oviducal
eggs were. — June: 5.07 mm, 5.15 mm (female, 44.2 mm); July: 3-62
mm, 3.77 mm (female, 45.3 mm); August: 3.24 mm, 3.33 mm (fe-
male, 47.2 mm). The smallest maximum follicle diameters for sex-
ually mature females were. — June: 1.42 mm, 1.62 mm (female, 39.2
mm); July: 1.43 mm, 1.63 mm (female, 43.6 mm); August: 0.80
mm, 1.59 mm (female, 37.2 mm). The largest maximum follicle
diameter for females less than 35 mm snout-vent were. — June: no
data; July: 0.34 mm, 0.38 mm (female, 28.7 mm); August: 0.19
mm (female, 29.2 mm). The smallest follicle diameter maxima

22 Tulane Studies in Zoology Vol. 1

for females of this group were. — June: no data; July: 0.19 mm (fe-
male, 22.5 mm); August: 0.19 mm (female, 29.2 mm). The aver-
age maximum follicle diameters, exclusive of females with oviducal
eggs, for sexually mature females were. — June: 2.51 mm; July: 300
mm; August: 1.97 mm.

From the above data it is concluded that the egg laying season ex-
tends from March or April at least through August, reaching its
peak in May, June, and July. Cagle (1948b) reports a similar pro-
longed period for Anolis in Louisiana. As the largest follicles ob-
served were 5.15 mm and 5.07 mm in diameter, it is assumed that
ovulation occurs when follicles approach this diameter. The absence
of follicles of 1 mm or more in diameter during the months of Oc-
tober and December (except for two individuals, as noted above),
but their presence during other months of the year, is presumptive
evidence that such follicles are either ovulated or resorbed during
the season of their occurrence. The fact that no females less than 35
mm snout-vent possessed follicles of this diameter at any time dur-
ing the year, and especially during the breeding season, is the basis
for using the criterion of follicles equal to or greater than 1 mm in
diameter as evidence of sexual maturity. That occassional females
may contain oviducal eggs and follicles of 2 mm and more in diameter
suggests the possibility of such females having two broods per season.
Futher study is necessary to establish this point. Atsatt (1953: 59)
cites several distinct clutches in one season for the dwarf chameleon
Microsaura pumila pumila ( Daudin ) .

Hatchlings are first in evidence in June (figs. 11, 12). These
individuals are in the 16-19 mm snout- vent size group. Davis (1945:
116) gives 21 mm (total length minus tail length) as the average
snout-vent length of three hatchlings. All individuals of this size
group contained remnants of the yolk sac. The absence of this size
group in other months of the breeding season does not support the
contention of two broods per season. There absence probably reflects
the bias of collecting methods rather than the activities of the lizards.

Examination of the oviducal eggs disclosed the presence of an
advanced embryo in each of 30 eggs of 10 females collected in
March, June, July, and August. The embryos ranged from 1.54 mm
to 3.10 mm in length. All were in the somite stage with brain and
tail flexures. Weekes (1927a; 1927b) reports ovoviviparity for Old
World species of Lygosomas

GROWTH

Conclusive data for determination of growth rate are not available
(figs. 15, 16). An estimate of growth rate is necessary to postulate
age at which sexual maturity is attained. Except for a male in March
and one female each in January and April, all individuals collected
from January through May are of adult size (figs. 11, 12). From June
through August three size groups are present: hatchlings, sexually

No. 2

Johnson: Life History of Scincella

mature individuals, and an intermediate size group. This intermed-
iate size group may be explained by one or both of two causes.

8+.337X

7u — 7y — Ku — S — 5 — H — D — 75 — t — R — T

Months

rig. 15.

Correlation of Snout-Vent Lengths (Means

: Three Standard Errors) With Month! (Males).

Ju Jy Au S 0 N D Ja F H a N

Months

Fig. 16.

Corrtlatloa of Snout-Vent Lengths (Maana
t Three Standard Errors) With Months (Fsmaloa).

As mentioned above, no literature records of eggs collected in the
field (or laboratory) prior to May were found. Inferring a minimum
of four weeks for incubation (the interval from the date of earliest
field collected eggs and the appearance of hatchlings), the intermediate
size group is best explained as second brood or late single brood in-
dividuals of the previous season. These are immature yearlings.
That this intermediate size group constitutes early (prior to May or
June) hatchlings of the current season is a second less plausible ex-
planation. In either event, most individuals from 24-25 mm snout-
vent present in June (?), July, and August would not contribute
to the current breeding population.

The distinct size groups present from June through October merge
by November and constitute a single group with wide range of snout-
vent length measurements. The individuals in the lower snout-vent
length classes would be composed primarily of the current season's
hatchlings. Late hatchers of the previous season cannot be dis-
tinguished. By January the limits of the snout-vent length measure-
ments have been decreased as a result of the increase to mature snout-
vent length size of the past season's hatchlings. On the basis of these
data, it is assumed that sexual maturity may be attained by the season
following hatching if that occurs in May through August. The

24 Tulane Studies in Zoology Vol. 1

late hatchers (subsequent to August) may not contribute to the
breeding population until very late the following season or even the
second season following hatching. This problem cannot be resolved
without further and more intensive study of growth rates.

Differential growth between males and females is evident. The
most apparent difference is a longer axilla-groin length relative to
snout- vent length for females (fig. 6). Females also attain a larger
snout-vent length than males. In both males and females there is
allometric growth of axilla-groin length and length of limbs. The
adpressed limbs fail to meet in most males more than 35 mm snout-
vent. Three males 41.1 mm, 44.9 mm, and 45.8 mm had overlap of
the adpressed limbs. Among the females there were none in excess
of 39 mm snout-vent having overlap of adpressed limbs. Similar
allometric growth occurs in Mexican forms of this genus (Taylor,
1937).

ACKNOWLEDGEMENTS

I am grateful to Dr. Fred R. Cagle for his help and criticisms dur-
ing the progress of this study. Drs. E. S. Hathaway, George H. Bick,
and George H. Penn also receive my expression of gratitude as do
the fellow students who assisted and encouraged me.

I wish to thank Drs. W. D. Stull, C. O. Berg, and W. F. Hahnert,
Department of Zoology, Ohio Wesleyan University, for their crit-
icisms of the manuscript.

SUMMARY

1. Four hundred and sixty-two specimens of Scincella later ale
(Say) were examined to determine size at sexual maturity, seasonal
reproductive cycle, and reproductive potential.

2. A series of 32 males and 37 females (Tulane 14108) over
35 mm snout-vent lengths was examined to determine sexual di-
morphism in selected characters. Sexual dimorphism is present as
regards axilla-groin/snout-vent length, snout-vent/tail length, and
dorsal scale row counts, with females having larger axilla-groin/snout-
vent length ratio and dorsal scale row counts and males having a
larger snout-vent/tail length ratio. No dimorphism exists for mid-
body scale row counts or pigmentation.

3. Juveniles were observed to show sexual dimorphism only
with respect to dorsal scale row counts as series Tulane 14108 above.
Dimorphism with respect to axilla-groin/snout- vent length begins
at about 24 mm snout-vent.

4. A positive correlation between axilla-groin and snout-vent
lengths exists for both sexes. Males and females differ significantly
in this character.

5. Males and females exhibit allometric growth of the axilla-
groin length and limb lengths. Adpressed limbs of males less than
35 mm snout-vent and females less that 39 mm snout-vent overlap.

No. 2 Johnson: Life History of Scincella 25

6. The scales on the sides of the neck, base of the tail, in the
axilla and groin, and on the posterior surfaces of the limbs are keeled.
Occasional individuals may have keels on the dorsal scales as well as
the above areas. This is especially true of juveniles. No sexual di-
morphism exists as regards these keels.

7. No geographic variation exists for the Louisiana population
as regards certain selected morphological characters.

8. Males are sexually mature at 35 mm snout- vent length. Sper-
matozoa are present in the testes and vas deferens from January
through August.

9. Females are sexually mature at a snout-vent length of 35 mm.

10. Ovarian follicles 1 mm or more in diameter are present from
December through August. These follicles are assumed to indicate
the extent of the breeding season for females. The concurrence of
oviducal eggs and follicles of 2 mm or more in diameter is indicative
of the possibility of two broods per season. Only females of 44 mm
or more in snout-vent length show this potentiality. Oviducal eggs
are present from March through August.

11. The average number of oviducal eggs per female is 3-3 —.05,
the range from one to five. Eggs were found in the right oviduct
only, but not in the left oviduct only. The average number of eggs
in the right oviduct, 1.9— .06, is significantly different from the
average of the left oviduct, 1.5— .05.

12. Hatchlings are those individuals between 16-19 mm snout-
vent. Individuals hatched from May to August may reach maturity
by the spring following hatching. Late hatched individuals (subse-
quent to August) may not reach maturity until very late the follow-
ing season or even until the second season following hatching.

13. Embryos advanced to the somite stage with brain and tail
flexures were observed in 30 oviducal eggs from ten females col-
lected in March, June, July, and August.

14. Field studies of egg laying activities, hatching dates, and
growth rates are needed to corroborate the assumptions concerning
these activities based on laboratory studies.

26 Tulane Studies in Zoology Vol. 1

REFERENCES CITED

Atsatt, S. R. 1953. Storage of sperm in the female chameleon Mi-
crosaura pumila pumila. Copeia, 1953: 59.

Blanchard, Frank N. 1921. A revision of the king snakes: genus
Lampropeltis. U. S. Nat. Mus., Bull. 114: 1-260.

Blount, R. F. 1929. Seasonal cycle of the intersticial cells in the
testes of the horned toad (Phrynosoma solar e) . Jour. Morph.
and Physiol, 48: 317-343.

Breckenridge, W. J. 1943. The life history of the blackbanded
skink, Eumeces s. septentrionalis (Baird). Amer. Midi. Nat.,
29: 591-605.

Cagle, Fred R. 1948a. Sexual maturity in the male turtle, Pseudemys
scripta troostii. Copeia, 1948: 108-111.

1948b. A population of Carolina anol'e. Chicago Acad. Sci., Nat.
Hist. Misc., No. 15 : 1-5.

Carr, Archie F. 1940. A contribution to the herpetology of Florida.
Univ. Fla. Publ, Biol. Sci. Ser., 3 : 1-118.

Cazier, Mont A. and Annette L. Bacon. 1949. Introduction to quan-
titative systematics. Bull. Amer. Mus. Nat. Hist., 93 : 349-388.

Cieslak, Edwin S. 1945. Relations between reproductive cycle and
the pituitary gland in the snake Thamnophis radix. Physiol.
Zool, 18: 299-329.

Cook, Fannye, A. 1943. Alligators and lizards of Mississippi. Surv.
Bull., Miss. State Game and Fish Comm., pp. 1-20.

Davis, William B>. 1945. The hatchling of Leiolopisma laterale. Co-
peia, 1945: 115-116.

Klauber, Lawrence M. 1937. A statistical study of the rattlesnake.
Occ. Pap., San Diego Soc. Nat. Hist., No. 3 : 2-56.

1943. Tail-length differences in snakes with notes on sexual
dimorphism and the coefficient of divergence. Bull. Zool. Soc.
San Diego, No. 18 : 5-60.

Lewis, Thomas Howard. 1951. The biology of Leiolopisma laterale
(Say). Amer. Midi. Nat., 45: 232-240.

Mayr, Ernst. 1942. Systematics and the Origin of Species. Columbia
University Press, New York, pp. 3-334.

Mittleman, M. B. 1947. American Caudata. I. Geographic variation
in Manculus quadridigitatus. Herpetologica, 3: 209-224.

1950. The generic status of Scincus lateralis Say, 1825. Ibid.,
6: 17-20.

Ortenburger, Arthur Irving. 1928. The whipsnakes and racers,
genera Masticophis and Coluber. Univ. Mich. Studies, Mem. Univ.
Mich. Mus., 1 : 1-247.

Peatman, John Gray. 1947. Descriptive and Sampling Statistics.
Harper and Brothers, New York, pp. 3-577.

Reynolds, W. E. 1943. The normal seasonal cycle in the male Eu-
meces fasciatus together with some observations on the effects
of castration and hormone administration. Jour. Morph., 72:
331-377.

Simpson, George G. and Anne Roe. 1939. Quantitative Zoology.
McGraw-Hill Book Company, Inc., New York, pp. 1-414.

No. 2 Johnson: Life History of Scincella 27

Smith, Hobart M. 1946. Handbook of Lizards of United States and
Canada. Comstock Publishing- Company, Ithaca, pp. 1-557.

Snedecor, George W. 1946. Statistical Methods. Iowa State College
Press, Ames, pp. 1-485.

Taylor, Edward S. 1937. Two new lizards of the genus Leiolopisma
from Mexico, with comments on another Mexican species. Copeia,
1937: 5-11.

Viosca, Percy, Jr. 1933. Louisiana Out-of-Doors. Author, New Or-
leans, pp. 1-187.

Weekes, H. Claire. 1927a. A note on reproductive phenomena in
some lizards. Proc. Linn. Soc. New South Wales, 52: 25-38.

1927b. Placentation and other phenomena in the scincid lizard
Lygosoma (Hinulia) quoyi. Ibid., 52: 499-554

INI

-j

Jim ^®(DiL(DOT

Volume 1, Number 3

July 28, 1953

AN OUTLINE FOR THE STUDY OF A REPTILE
LIFE HISTORY

FRED R. CAGLE

DEPARTMENT OF ZOOLOGY, THE TULANE UNIVERSITY OF LOUISIANA,

NEW ORLEANS.

Ml CO&P. ZSSL

library

AUG 2 4 1953

HARVARD
UNIVERSITY

TULANE UNIVERSITY
NEW ORLEANS

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need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

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Reptiles, American Museum of Natural History, New-
York. N. Y.

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History Museum, Chicago, Illinois.

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mittances should be made payable to Tulane University.

Price for this number: $0.75.

George Henry Penn, Editor,
c/o Department of Zoology,
Tulane University of Louisiana,
New Orleans, U. S. A.

AN OUTLINE FOR THE STUDY OF A REPTILE
LIFE HISTORY

FRED R. CAGLE

Department of Zoology, The Tulane University of Louisiana,

New Orleans.

The renewed interest of biologists in natural populations, the de-
velopment of the new systematics, population genetics, biodemography,
and biosociology, i.e. of bionomics, or ecology in the broadest sense,
has brought demands for detailed information on life histories of
animals. The herpetologist is thus obligated to reexamine many of
the standards and customary procedures in natural history. Renewed
emphasis is also being placed on the organization of the knowledge
comprised in natural history. Investigations in autecology (the
ecology of the individual or of the individual species) are aimed
toward the development of significant generalizations and principles.
The modern systematist should not and must not divorce his thinking
from ecology; the ecologist must not ignore systematics. While con-
demning the ecologist for ignoring systematics, the systematist has
often been guilty of ignoring ecology. There is an urgent need for
men thoroughly trained in the techniques of both ecology and sys-
tematics, men who can accelerate the trend toward a blending of
these fields.

The investigator interested in reptilian populations finds but few
studies of the bionomics of reptiles that meet critical standards. He
finds an assortment of fragmentary facts that are difficult if not im-
possible to integrate, and often immediately require the test of repe-
tition. It may be pointed out that repetition of field observations
in a critical spirit may be fully the equivalent of experimental test.
There seems to be a need for a statement of minimum requirements
of information basic to the formulation of suggestions for a systema-
tic approach to research on natural populations.

Perhaps the best test of significance of an observation in "natural
history" is one similar to the test for the adequate description of a
species or subspecies. Does the observation reflect an attribute of a
given population? Is it reported in such a fashion that it may be
integrated with other observations to state such an attribute? Or,
has the research merely reported an aberrant or extreme behavior
pattern, an anomalous situation, or such fragmentary data that it
fails to express any particular fact as an attribute of a population?

What information is required in a definitive life history investi-
gation? Ideally? What, practically, can the investigator hope to
contribute? In what areas of biology may his data be applicable?
What are the prevalent fallacies in life history analyses? Some at-
tempt is made here to indicate limits and to designate specifically
the obligation of the student interested in natural populations of

32 Tulane Studies in Zoology Vol. 1

reptiles. Many of the techniques developed by the ichthyologist,
mammalogist and ornithologist may well be utilized by the herpetolo-
gist. There is an urgent need for the development of new techniques
of research and for new applications of old ones. The herpetologist
does not have the equivalent of the procedure of the ichthyologist
for determining the age and growth of an individual by the exami-
nation of the growth lines of scales; nor does he have the trapping
procedure for systematic sampling as used by the mammalogist, or
the activity recording techniques developed by the ornithologist.
Equivalents of these techniques are among our greatest needs. All
of the needs for herpetological investigation cannot be enumerated
in this paper, nor can all of the techniques developed in other fields
be mentioned, but the bibliography is intended to provide suggestions
and a key to the vast literature.

The herpetologist concerned with the study of a single form cannot
hope to explore intensively all of the questions presented in the fol-
lowing discussion, but his awareness of the problems and of the
need for information will permit him to make observations that
otherwise might not be recorded. A serious report on a life history
should be the result of a planned, long term research project. Such
a report will integrate all of the minor elements of the topic to be
gleaned from the literature with the more directed accumulation of
planned observations of the author. The investigator, through proper
planning, may maintain several such research programs. The outline
that follows is purposely elaborated; and it includes much detail that
may appear unnecessary to my colleagues; but it is directed to the
students of the future who may be approaching similar problems
from both the ecological and the systematic side. An early and com-
parable effort to systematize studies in life histories that has had a
long usefulness is the summary of Walter P. Taylor (1919). Fitch
(1949) presents many valuable suggestions for the student interested
in natural history.

This paper is the outgrowth of an outline for the study of a reptile
life history prepared originally under the direction of Dr. Norman E.
Hartweg, University of Michigan. Its development has been en-
couraged by the critical and generous comments of Mr. Karl P.
Schmidt, Chicago Natural History Museum. I am indebted to Mr.
Roger Conant, Philadelphia Zoological Garden and Mr. Arthur
Loveridge, Harvard University for their suggestions. The group of
graduate students in herpetology at Tulane University has been a
constant source of stimulating challenge in the preparation of this
report. Mr. A. H. Chaney, Mr. Robert Gordon, Mr. Paul Anderson
and Mr. Richard Johnson have been of particular aid.

Studies contributing to concepts expressed in this review were
aided by a grant from the National Science Foundation.

I. What are the morphologic characteristics of the population to
be studied? Is action taken to insure that the data reported are

No. 3 Cagle: Outline for Reptile Life History 33

obtained only from individuals of the genus, species or sub-
species intended to be studied?

A. What is the taxonomic status of the population? What are
the diagnostic features? How are these related to the formal
description of the species or subspecies? Are these sharply
or only obscurely characterized?

These data are of extreme significance. Excellent infor-
mation must often be discarded by subsequent workers
because the author has failed to indicate clearly the taxo-
nomic characteristics of the population studied. Descrip-
tion must thus be such that any investigator can recognize
the population regardless of changes in nomenclature.

B. What other names have been attached to this population?

C. What samples of the population were collected and pre-
served and where are they deposited? Museum numbers?

A representative series supporting the description given
must be collected and deposited in a suitable museum col-
lection. Failure to do this is almost characteristic of
ecological investigations, yet the conclusion submitted
are often not acceptable because of questionable identifi-
cation of the material on which they are based.

D. What variation is observed in the individuals composing
the population? Of what is this variation a reflection?
Precise analysis and explanation of individual variation is
an obligation. Dice (1952) points out that few museums
have adequate storage or curatorial facilities to retain the
large number of specimens necessary for the analysis of
variation in local populations. The investigator must often
utilize materials that cannot be available to future workers;
his responsibility is thus multiplied. Through such studies
associated with field investigations we may hope to accumu-
late the data basic to systematic studies at the intraspecies
level.

1. What changes in color intensity, in pattern, or mor-
phology occur from birth to old age? Are there any
correlated sex differences? How are these changes re-
lated to taxonomic investigations? The limited infor-
mation available on ontogenetic changes in "characters"
has resulted in much confusion in taxonomy. The
trend in herpetological research toward thorough analysis
of such changes promises the development of a basis
for substantial clarification of the status of many forms
(Oliver, 1951).

2. Is the variation correlated with differences in the ex-
ternal environment?

With a gradient in the external environment? Is the

34 Tulane Studies in Zoology Vol. 1

the variation due to differences in genotypes or does it
reflect the responses of a specific genotype to different
environments?

Investigations often query the status of the variation
described but do not perform the simplest of experi-
ments aimed at evaluating the genotypic flexibility
of the organism studied. Some investigations suggest
that some of the characters considered to be of taxo-
nomic importance are merely phenotypic modifications
(Fox, 1948). It is essential to the systematist that
he determine whether variations are the result of
heredity or environment or both.

3. What are the ontogenetic changes in mass as expressed
by measurements or weight? What is the maximum size
attained? Sex differences?

Although absolute size is not an acceptable taxonomic
character for poikilothermic vertebrates, genetic dif-
ferences in potential natural longevity or growth
potentials may be reflected in differences in maximum
sizes between populations (Lagler and Applegate,
1943).
What procedures were used in mensuration? Weigh-
ing?

Care must be used to insure adequate mensuration
practices and to insure that the investigator clearly
reports his procedures (Simpson and Roe, 1939;
Cazier and Bacon, 1949). Much confusion has been
caused by misunderstandings resulting from failure
to specify the methods followed. The significance of
the limits of error in such data should be borne in
mind. When measurements are accurate only to milli-
meters, proportions calculated to three decimal places
give a false aspect of accuracy of the data.

4. What are the principle differential growth changes in
each sex? How are these changes related to the major
phases of the life history?

Failure of the systematist to recognize the presence
of differential growth has led to the erroneous use of
proportions. If detailed quantitative studies cannot
be made, the investigator should, as a minimum, desig-
nate the gross changes in proportion. This is a par-
ticularly acute problem in poikilothermic vertebrates
(Hersch, 1941).

II. What is the geographic range?

The range should be expressed first in terms of museum
specimens or records of authorities. All questionable records

No. 3 Cagle: Outline for Reptile Life History 35

should be deleted. The range definition should indicate the
distribution of existing populations (Grobman, 1950). These
data may then, in connection with other information, form
the basis for the statement of a supposed "true range".

A. What are the factors limiting the range?

These must be considered in terms of the ecological data
assembled during the progress of the investigation with
particular reference to the total knowledge of the eco-
logical valence of the animal and possible barriers to dis-
persal (Darlington, 1948; Cowles and Bogert, 1944; Dice,
1952). It is especially important to note that the limit-
ing factors may be entirely different on the different
borders of the range of a species (Schmidt, 1950).

B. What physiographic and climatic factors are characteristic
of the range?

1. What are the annual temperature and rainfall cycles?

2. What are the mean annual, minimum and maximum
temperatures in the warmest and coldest parts of the
range?

Whenever feasible, temperature and rainfall data col-
lected by the investigator in the areas of intensive
study should be utilized. Of necessity, the investi-
gator must often use meterological and climatological
temperatures, but their interpretation should be based
on the data of the researcher (Baum, 1950).

3. Does temperature summation (heat summation) affect
the distribution of the species investigated?

C. What is the principal habitat? Marginal habitat?

1. Are microclimates of significance? throughout the
range? at the periphery of the range? (Geiger, 1950;
Diem, 1951).

2. What vegetational types characterize the habitat?

3. Do size or age groups tend to occupy different habitats?

4. Does the animal have an innate habitat recognition
mechanism? (Svardson, 1949; Tinbergen, 1948).

HI. What is the age and sex composition of a local population?

A. What annual changes occur in the composition of a local
population?

1. What is the sex ratio in mature individuals during the
breeding season? How does this change during a single
year?

Sex ratios are often reported without reference to ma-
turity or to the breeding season although radical
changes do occur in some reptile populations. Esti-

36 Tulane Studies in Zoology Vol. 1

mates of the relation of sex ratios to natality should
be based only on the relative frequency of mature in-
dividuals (Forbes, 1940; Cagle, 1948).
Sex identification is frequently reported without refer-
ence to the criteria used. What are these criteria?
Secondary sex characters? Gonad condition? If dis-
section, on what basis was sex determined?

2. What is the sex ratio in juveniles? At birth? In pro-
gressive age groups?

3. What annual changes occur in the ratio of juveniles to
adults? What is tne potential contribution from "young
of the year" to the adult segment of the population?

4. Can an ecological life table be constructed?

The difficulty of determining mortality rates in most
reptiles forbids the successful completion of such
tables yet an attempt to collect data basic to the esti-
mation of survivorship curves should be made (Dee-
vey, 1947).

5. What are the major predators? Is predation pressure
a significant factor in annual and long term cyclic
changes? What is the relation of loss from predation to
population density? (Errington, 1946).

B. What long-term cyclic changes occur in the composition of
the local population? What is the cause of such cycles?
Is exhaustion of the adreno-pituitary system a factor as has
been demonstrated for some mammal populations? (Chris-
tian, 1950; Elton, 1942).

C. Do local populations differ in composition? If so, what is
the basis of such differences?

Adequate local sampling provides a basis for obtaining
answers to such questions. It has been demonstrated that
substantial differences may be present in the compositions
of local populations. Comparison of population samples
must be tempered with an awareness of the difficulties
of obtaining such samples. Series of specimens preserved
in museum collections are rarely unbiased samples of
natural populations. The student should note particularly
those few long-term studies in local areas (De Haas,
1941).

D. Does the individual animal or the mated pair occupy a
home range (or activity range as defined by Carpenter,
1952). Territory?

1. What is the size of the home range and of the territory?

a. What features of the habitat may modify the size?

No. 3 Cagle: Outline for Reptile Life History 37

b. What is the relation of the size of the territory or
home range to density?

c. Does the individual have homing ability? If so,
what are the mechanisms involved in orientation?

The recovery of marked individuals in short-term
and long-term studies will provide information on
these questions. A wide variety of methods
have been used for the marking of reptiles: metal
bands or plates (Wickham, 1922); scale clipping
(Blanchard and Finster, 1933; Conant, 1948; Fitch,
1949); plate notching (Cagle, 1939); tattooing
(Woodbury, 1948); branding (Woodbury and
Hardy, 1948); painting (Cagle, 1946). Trapping
and other special collecting procedures are de-
scribed by Dargan and Stickel (1949), Lagler
(1943a). The calculation of size of home range
from trapping results is discussed by Hayne
(1949) and Stickel (1950). Stickel and Cope
(1947) summarize information on home ranges.

Schaefer (1941), Bailey (1952) and Leslie (1952)
discuss the estimation of size of animal popula-
tions by marking experiments.

The multiplicity of problems involved in animal
orientation are ably discussed by Fraenkel and
Gunn (1940).

2. Is the territory selected by the male, female or both?

Do both sexes participate in its defense?

a. What are the characteristic behavior patterns used
in defense of territory?

Lowe and Norris (1950) summarize the reports
of aggressive behavior in snakes.

b. What is the chief stimulus to maintenance of ter-
ritory?

c. Is the territory maintained throughout the year or
or only during short periods?

Nice (1941) presents a classification of the types
of territoriality.

IV. What is the density of the population?

There should be more than a vague estimate of density ex-
pressed as rare, common or abundant. The objective should
be to gain a measure of the number of individuals in a given
area expressed in terms clearly defined by the investigator.
The use of the concepts of abundance, and relative apparent
abundance as suggested by Marr (1951) is recommended.
The method selected for this determination of abundance

38 Tulane Studies in Zoology Vol. 1

must rest on the knowledge of the ecological requirements
of the individual. Kendeigh (1944) provides a suggestive
review of the procedures for measurement of bird population.
Andrushko (1936) suggests techniques suitable for some
species. A summary of methods is presented by Thomas
Park (1950). Information of particular value in estimating
populations from recovery of marked specimens is given by
Ricker (1948), Jackson (1939) and Bailey (1952). This
procedure has been applied to reptiles by several authors
(Cagle, 1950; Fitch, 1949; Stickel, 1950).

A. What is the relation of density to the questions posed in
sections I, D and III A to D (Blair, 1951)?

B. What is the relation of density of the form studied to that
of other reptiles inhabiting the area? (Fitch, 1949; Cagle,
1950; Cagle and Chaney, 1950).

V. What is the potential reproductive capacity? What is the rela-
tion to realized reproductive performance? What are the best
measures of natality?

A. At what age and/or size does the animal become sexually
mature?

1. When are the secondary sex characters developed?
What is the relation of time of their appearance to the
potentiality of sexual functioning? (Regamey, 1935)

2. What cyclic changes occur in secondary sex character-
istics?

3. What is the relation of age of attainment of maturity
to the annual reproductive cycle?

Investigators often fail to indicate what they mean
by sexual maturity. Care must be exercised that the
criteria for maturity are defined. In reptiles these
may concern the presence of oviducal eggs in females,
of corpora albicantia, of ovarian follicles of a speci-
fied size or ovaries of a specified weight or volume
(Altland, 1951). In males a specific stage of sperm-
atogenesis, a specified testicle weight or volume in
relation to an indication of total body mass or the
presence of motile sperm may be useful (Cieslak,
1945; Cagle, 1944; Risley, 1938; Fox, 1952). No
adequate techniques are available for determining the
age of an individual reptile. The procedures used by
Bryuzgin (1939) should be further explored. Bryuz-
gin concluded that rings discernible in cleared skull
bones of snakes could be used to determine age.

B. What is the total period of reproductive activity in the life
of an animal?

No. 3 Cagle: Outline for Reptile Life History 39

1. Does the annual reproductive potential remain the same,
decrease or increase with age?

2. When does senility occur?

3. What is the ecological longevity?

C. What is the annual realized reproductive performance?

1. What is the annual period of reproductive activity in
females? in males? What is the relation of this period
to the total annual activity cycle?

Baker (1947) discusses the causes of breeding seasons.
Vols^e (1944) describes seasonal fluctuations in the
reproductive system. Kendeigh (1941) summarizes
information on the relation of length of day to gonad
development. This period is usually considered as
that period in which the females are "carrying" young
or are laying eggs. Much confusion has resulted from
failure to delimit this period. Thus it may be stated
that a female having eggs in the oviduct was col-
lected on a given date. Yet this is not clearly in-
dicative of the time when eggs may be deposited.
Each investigator should insure preciseness of defi-
nition. Writers frequently use the presence or ab-
sence of oviducal eggs to delimit the season but this
can lead to potential errors if not weighed properly.
Eggs may be retained in the oviducts for long periods
(Cagle and Tihen, 1948).

2. What correlation is there between courtship or copula-
tion and ovulation? What is the significance of the sex
ratio and population density in relation to annual real-
ized reproductive performance?

These are little-explored areas in herpetology yet im-
portant ones if we are to arrive at an understanding
of those factors controlling changes in reptile popu-
lations. The fact that some reptile females may bear
young or deposit fertile eggs after as long as eight
years after copulation suggests that unbalanced sex
ratios may be of but scant consequence. The unveri-
fied yet not disproven statement that single or suc-
cessive copulations are essential to stimulate ovulation
indicates the importance of a favorable sex ratio. The
work of Darling (1938), Vogt (1942), Errington
(1946) and others has suggested that population
density may markedly affect breeding success.

a. What is the pattern of courtship?

Exploration of the courtship patterns with em-
phasis on interspecies differences promises to
yield much of value in explaining the develop-

40 Tulane Studies in Zoology Vol- 1

ment of physiological isolation. Noble and Brad-
ley (1933) furnish many suggestions for pro-
cedure and interpretation. Cagle (1950) de-
scribes differences in the courtship pattern be-
tween two species of the genus Pseudemys. Davis
(1936) summarizes the literature for snakes;
Gloyd (1947) suggests additional problems;
Greenberg (1945) summarizes the knowledge of
courtship in the family Iguanidae*

(1) How does it differ from that of related forms?

(2) What advantages in reproduction are provided
by the courtship pattern?

(3) What selective factors function in courtship?

(4) What secondary sex characters are of most
significance in courtship?

(5) What senses are involved in courtship? (Noble,
1937)

b. What is the relation of courtship drives to aggre-
gation? (Finneran, 1949).

c. When do ovulation and fertilization occur?

(1) What is the fertilization rate? The relation of
successful courtship and copulation to fertiliza-
tion rate?

(2) Is copulation essential to ovulation? to egg
depositions? (Woodward, 1933).

3. How many groups of young (eggs) are produced each
year?

This question must usually be answered by the exami-
nation of ovaries from chronological samples taken
during the breeding season so that progressive changes
in number and size of ovarian follicles or total volume
or weight may be reported. Too, examinations of
the ovaries of females at the end of the reproductive
period may yield counts of ovulation points (corpus
luteum or corpus albicans) (Samuel, 1952).

4. How many young (eggs) are produced in each group?

Some investigators have depended solely upon counts
of oviducal eggs or of eggs found in nests. Both pro-
cedures are subject to substantial error as the worker
can but rarely be confident that no eggs have been
previously deposited, that ovluation is completed or
that two or more females have not utilized the same
nest. Counts of ovulation points are usually more
acceptable. Certainly the typical extreme variation
in number of eggs and young produced emphasizes
that little significance may be attached to many of

No. 3 Cagle: Outline for Reptile Life History 41

the literature reports of the number of young in single
females or nests. Counts of young present in the
uteri of viviparous or ovoviviparous forms possibly
provide the most reliable criteria of clutch size. (The
terms, viviparous and ovoviviparous, have been used
in varied ways in herpetological literature. It is sug-
gested that the term, ovoviviparous, be restricted to
describe a situation in which the developing young
gains no sustenance from the female),
a. Is there a correlation between reproductive capacity
and size or age? How is this related to estimates
of natality in local population?

The large difference in reproductive capacity be-
tween small and large females make it exceedingly
difficult to utilize much of the published data on
reproductive capacity as bases for estimates of
natality.

VI. What are the major factors controlling the relation of the
number of surviving young to the number of eggs or young
produced by females?

A. What are the characteristics of the egg at deposition?

1. How do the eggs vary in size, volume and weight in each
clutch?

The irregular shape of most reptile eggs reduces the
value of measurements of length or width reported
without volumes (Lynn and Brand, 1945).

2. Is there any correlation in size and/or weight and size
of female?

3. What changes occur in size and weight of eggs during
incubation?

The weight and volume of eggs change much and
irregularly with age and the environment. Cunning-
ham and Hurwitz (1936) reported that eggs increased
as much as 60% in weight during incubation. Data
on reptile eggs are of little value unless they are ac-
companied by statements as to their age and condi-
tions under which they were incubated. The statis-
tical treatment (Edgren, 1949) does not remedy this
discrepancy.

4. In what stage of development is the egg at deposition?

a. Does this stage. of development vary with the time
eggs are retained in the oviducts? If so, how does
this influence the incubation period?

b. How is the stage of development related to the egg
size and weight?

42 Tulane Studies in Zoology Vol. 1

B. Where and in what manner are eggs deposited?

1. Is a nest constructed?

a. What factors determine the nest site?

b. How is the nest constructed?

c. What is the relation of choice of nest site and con-
struction to potential survival of young?

d. What is the behavior pattern of the female construct-
ing a nest? What features are of survival import-
ance?

e. Does the female use the same nesting site for subse-
quent clutches? in subsequent years?

2. Does the female remain with the eggs? return to them?
What is the relation of female behavior to survival
potential of young? of the females?

a. Does the female "defend" the eggs?

b. Does the female contribute "heat" to incubation?

These questions cannot be answered on the basis
of single observations. Behavior of reptiles is
sufficiently variable that repeated observations are
essential to description of behavior patterns. In
most situations the investigator can gain but re-
stricted field data on these questions and is com-
pelled to study captive specimens as a basis for
evaluation of field-collected data (Noble and
Mason, 1933).

C. What factors determine incubation rates?

1. What is the period of incubation? in field nests? in
the laboratory?

a. What is the relation of temperature levels or changes
to incubation time? of degree-hour to incubation
time? (Cunningham, 1939).

b. Are differences in incubation time between clutches
of eggs related to egg-deposition (sequence in ovi-
ducts; time of retention in oviducts; quality of shell
deposited)? Observers frequently do not state in-
cubation periods in degree-hours and do not provide
their criteria for "hatching". The extreme difficulty
of evaluating much of the published material makes
it unavailable for coherent treatment.

It is not usually possible to observe the deposition
of reptile eggs and the incubation period must be
expressed as the interval between the laying of the
last egg and the hatching of the last egg. This pro-
cedure is usually followed in reporting the incubation
time of bird eggs (Skutch, 1950). Although rep-

No. 3 Cagle: Outline for Reptile Life History 43

tiles typically deposit an entire brood over a short
period as compared with birds, the total time re-
quired to deposit a brood is often significant in
relation to the incubation period. The incubation
time should be expressed in terms of days and hours
or degree-hours.

2. How sensitive are eggs to low or high temperature dur-
ing the incubation period? What extremes are the eggs
subjected to in the typical nest site? Potential mortality?

3. How do the hatchlings escape from the egg? What is
the function of the caruncle? What mortality is involved
in the process of hatching?

D. Does the female develop any particular behavior traits as-
sociated with gestation?

1. What is the period of gestation? (Bragdon, 1951).

2. What are the principle causes of mortality during em-
bryonic development?

3. Does the female tend to select a particular type of site
for the birth of the young? Relation of such selection
to potential survival?

4. Describe the birth of the young.

VII. What are the characteristics of the young? Are there any typical
behavior traits? What is the relation of the behavior pattern
to survival? to growth?

A. What advantageous resources in morphology, physiology, be-
havior patterns do the young adults possess? (Daniel and
Smith, 1947).

1. What is the amount of yolk retained? Is it utilized as
a source of nourishment? How long and under what
conditions will it serve to support the young?

2. How long do the young remain in the nest or with the
female? What factors influence the length of this
period? May young overwinter in the nest? Remain
with the female for prolonged periods? What relation
may this bear to survival potentialities?

B. What are the major hazards to which the young are exposed
immediately after leaving the nest or the female?

VIII. What are the characteristics of the groivth curve of individuals
of the local population?

A. What is the length of the growing season?

1. What are the factors serving to delimit the growing
season? Availability of food? Changes in environ-
mental temperature? Cyclic changes independent of
temperature?

44 Tulane Studies in Zoology Vol. 1

Various procedures have been attempted for determin-
ing the limits of the growing season. The actual
observation of initiation and cessation of growth
through study of seasonal samples is best but such
observations are difficult to obtain. The correlation
of formation of growth rings in turtles with season
has been attempted (Cagle, 1946). Too, once the
minimum and maximum effective temperatures of
a form are known they may be utilized to approxi-
mate the time of initiation or slowing of activity.
This does not, however, necessarily define the growing
season as it has been demonstrated that reptiles may
become quiescent during the winter although re-
tained at constant temperature. Evans and Hegre
(1940) have suggested that some genetic time factor,
distinct from the temperature factor, is operative in
reptiles.

2. What variations in length of growing season occur
within the area of investigation?

It is indicated by some researches that the time of
initiation or cessation of growth may vary significantly
from one local habitat or situation to another.

B. What is the annual increment (in that measure selected as
the best indicator of total change in mass) during each
season of the animal's life? What sex differences occur?

1. What are the factors influencing the rate of growth?
(size and/or age; senility, length of growing season,
social dominance).

2. What are the limits of variation in growth rates? How
does growth rate affect the attainment of maturity, na-
tality, mortality?

3. What age or size groups may be discerned? (Klau and
David, 1952).

4. Is growth potentially continuous throughout the life of
the individual?

C. What is the natural (ecological) longevity?

1. What longevity records are available from captive sepci-
mens?

2. What estimates of age may be made from the popula-
tion samples (Woodbury, 1951).

3. What are the characteristics of youth, maturity, old age?

IX. What is the annual cycle of activity and what factors exert
primary influence on the cycle? (Fitch and Glading, 1947;
Oliver, 1947).

A. What is the relation of the growing season to the period

No. 3 Cagle: Outline for Reptile Life History 45

(periods) of courtship, egg-deposition and birth of young?

B. What are the optimum, minimum and maximum effective
body temperatures?

C. What is the seasonal cycle in diel behavior (e.g., in bask-
ing) (Girons, 1947).

D. Are the animals quiescent during any period of the year?
Are aggregations formed?

1. What preparations are made for the period of quies-
cence?

2. Where do the animals spend the winter?

3. What environmental factors cause the initiation of
quiescence? renewed activity?

Bailey (1949) demonstrated that the plains garter-
snake, Thamnophis radix could endure temperatures
of approximately — 2°C. for a protracted period.

4. What is the composition (age groups, size groups, sex
ratios) of the winter aggregation?

5. What is the role of winter quiescence in limiting the
geographic distribution? (Bailey, 1948).

X. What is the diel cycle of activity?

A. What is the role of basking in the daily cycle?

1. What determines the time of basking, the length of the
period?

Sergeev (1939) reports a close relation between en-
vironmental temperature and the period of activity.
Benedict (1932) summarizes temperature relations in
reptiles.

2. What is the function of basking?

a. How is the period of basking related to rate of
increase or decrease of body temperature?

b. What is the characteristic behavior pattern in bask-
ing. How is this related to control of body tem-
perature? (Cowles and Bogert, 1944; Gunn, 1942;
Chernomordikov, 1943; Bogert, 1949).

B. Is feeding restricted to any particular part of the day? How
is the feeding behavior or length of the feeding period in-
fluenced by food availability?

C Are breeding activities (courtship; egg-deposition; birth of
young) restricted to any part of the day?

D. When does the peak of activity occur in the daily cycle?

E. How is the diel (Klauber, 1939) cycle modified by weather
changes, population density?

46 Tulane Studies in Zoology Vol. 1

Interspecies differences in the diel cycle of activity may
affect the entire life history. Exploration of the cycle
may yield the key to many of the problems presented
here. Noble (1946) presents valuable information on
such problems.

XI. What are the food habits? Their relation to growth and sur-
vival?

A. How does the animal obtain its food?

1. Can the animal pursue and catch actively moving prey?

2. What food preferences are exhibited in the field and
laboratory?

B. What are the principle foods? Relation to availability?

1. What is the relative importance of the food items?

2. How do feeding habits vary during the life of the ani-
mal?

3. Is there any seasonal variation in feeding habits?

Most studies of reptile food habits have reported a
high percentage of empty stomachs. It is thus es-
sential that the investigator utilize intestinal as well
as stomach contents. Too, the fecal material of
many reptiles may be used. Fitch and Twining
( 1946) emphasize the value of scats in the determi-
nation of the food habits of snakes. The scats of
lizards, particularly, are of great value in food analysis.
Carpenter (1952) obtained data on food habits of
snakes by forcing regurgitation. Lagler ( 1943b) re-
views the food habits of Michigan turtles.

C. Does the animal act as a controlling or limiting predator?

XII. Does this form exhibit any characteristic and genetically lim-
ited patterns of group behavior?

The study of behavior under undisturbed natural conditions
often yields startling information of basic importance to the
explanation of population problems (Svardson, 1949; Cal-
houn, 1950; Carpenter, 1950) and phylogeny (Bellairs and
Underwood, 1951). Few zoologists have developed the ability
to profit from the observation of field behavior patterns
(Emlen, 1950). Herpetologists, particularly have not utilized
this procedure.

A. Do aggregations occur? If so what are the stimuli and bind-
ing forces in aggregation? the function of the aggregation?
(Noble, 1936; Allee, 1931, 1951; Greenberg, 1943.)

B. Are social hierarchies present?

1. If dominance hierarchy is present, what is the relation

No. 3 Cagle: Outline for Reptile Life History 47

to territoriality, natality? (Evans, 1938, 1951; Greenberg,
1943).

2. How does the social hierarchy affect the migrating in-
dividual? the juvenile seeking a territory?

3. Does the social hierarchy influence growth and repro-
ductive potential? (Calhoun, 1950).

Such questions as these may be answered if some of
the methods of field ornithologists be adapted. The
use of blinds and optical equipment for observation
will yield much of value to the interpretation of
interactions. Observation towers were used to study
the behavior of turtles in Illinois (Cagle, 1944; 1950).
Excellent suggestions, many of which are of value to
the herpetologist, are presented by Emlen (1950).
The work of Evans (1938, 1951) is suggestive of
problems and procedures.

C. Are there typical defensive or offensive behavior patterns?
Bogert (1941) describes the "king-snake defense posture"
of rattlesnakes. Mertens (1946) summarizes reports of
such actions in reptiles.

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48 Tulane Studies in Zoology Vol. 1

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No. 3 Cagle: Outline for Reptile Life History 49

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1949a. Outline for ecological life history studies

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50 Tulane Studies in Zoology Vol. 1

1949b. Study of snake populations in central Cali-

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Fitch, Henry S. and Ben Glading 1947. A field study of a rattle-
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Fitch, Henry S. and H. Twining 1946. Feeding habits of the
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Forbes, Thomas R. 1940. A note on reptilian sex ratios. Copeia,
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Fox, Wade 1948. Effect of temperature on development of scutel-
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No. 3 Cagle: Outline for Reptile Life History 51

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^kasts sir® ©an

Volume 1, Number 4

August 15, 1953

A POPULATION OF HOLBROOK'S SALAMANDER, EURYCEA
LONGICAUDA GUTTOLINEATA (HOLBROOK).

ROBERT E. GORDON

DEPARTMENT OF ZOOLOGY, TULANE UNIVERSITY
OF LOUISIANA, NEW ORLEANS

MS. GOMP. 268L
LIBRARY

UJG 2 4 1953
HARVARD
UNIVERSITY

TULANE UNIVERSITY
NEW ORLEANS

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Coleman J. Goin, Associate Professor of Biology, University
of Florida, Gainesville, Florida.

Clifford H. Pope, Curator, Division of Reptiles and Amphibi-
ans, Chicago Natural History Museum, Chicago, Illinois.

Fred R. Cagle, Professor of Zoology, Tulane University of Lou-
isiana, New Orleans.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
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mittances should be made payable to Tulane University.

Price for this number $0.25.

George Henry Penn, Editor,

c/o Department of Zoology,
Tulane University of Louisiana,
New Orleans, U. S. A.

1AUG 2 4 1953
JHIVERSW I

A POPULATION OF HOLBROOK'S SALAMANDER, EURYCEA
LONG1CAUDA GUTTOL1NEATA (HOLBROOK).

ROBERT E. GORDON

Department of Zoology, Tulane University of Louisiana,

New Orleans.

Knowledge of the life cycle and general ecology of many of the
more abundant North American caudate amphibians is variously in-
complete or lacking. Holbrook's salamander, Eurycea longicauda gut-
tolineata is no exception. The literature records of this form are
based on a few recently metamorphosed individuals or larvae. No
samples have been available from which information relative to vari-
ation and population structure can be obtained. Two samples, col-
lected from the same population in 1950 and 1952, are deposited in
the collections of Tulane University (Tulane 13314 and 14905).

The author is indebted to Allan H. Chaney for information con-
cerning the 1950 sample, collected by the 1950 Tulane Field Crew,
and to Mr. Chaney and Robert G. Webb for aid in securing the 1952
sample. Both collections were taken during field trips made possible
by grants for summer research to Dr. Fred R. Cagle, to whom the
author owes much for opportunity to participate in the 1952 field
expedition and for valued criticism of this manuscript.

Both samples were taken approximately two miles south of Mari-
anna, Jackson Co., Florida on the Chipola River, the same locality
mentioned by Chaney and Smith (1950) and Tinkle (1952), al-
though the latter was misinformed as to the distance from Marianna.
Sixty-three individuals (Tulane 14905), collected in three man hours,
were taken on August 23, 1952 between 2:30 and 3:30 P.M. The
salamanders were collected in an irregularly shaped depression, ap-
proximately three-quarters of an acre in size, located in the river's
floodplain. Piles of debris and high water marks indicated previous
inundation, although no surface water was present. The vegetation
was predominately gum-cypress, with ironwood occupying the drier
edges of the area. The understory was sparse. A thick grass-herb
layer was encroaching toward the center from the drier sides of the
depression. The habitat fits Carr's (1940) description of the "Low
Hammock." Fallen timber and other surface litter was abundant.

The majority of the salamanders was collected from patches of
ground devoid of vegetation, but dense in surface debris and cypress
knees. Individuals were found under all sizes of debris; as many as
three were found under the same cover. The sky was overcast and
rain imminent. Several of the salamanders were active and could be
seen moving on the surface.

Associated species found under surface debris with Eurycea I. gut-
tolineata were Desmognathus fuscus fuscus, Microhyla carolinensis,
Eumeces fasciatus, and Diadophis punctatus punctatus.

The second sample, collected July 12, 1950, consists of eighty-one
specimens, approximately half of which (according to information
supplied by A. H. Chaney) were found in the area of the 1952
sample by Chaney. The other individuals were taken in adjacent
areas along the river.

Tulane Studies in Zoology

Vol. 1

Gonadal Development and Egg Deposition

The gonads of 144 individuals were examined macroscopically. The
gonads and accessary ducts of 56 specimens in the size range (snout-
vent) 21 to 39 mm (mean 34) are distinctly immature. The testes
and ducts contain no pigment. The compact ovaries contain many
small white ovocytes not enlarged with yolk material.

Eighty-eight specimens (varying in size from 43 to 61 mm, mean
52 mm) possess enlarged gonads and ducts when compared to the
above group. The testes and vasa deferentia are pigmented in vary-
ing degrees. The ovocytes appear to contain yolk material.

Forty -seven females (40 in the 1950 sample and 7 in the 1952
sample) constitute a size group falling within the range 46 to 61 mm,
average 53 mm. A correlation between condition of the oviduct,
ovary and body length is apparent (fig. 1). Twenty-four individuals
in the size range 46 to 54 mm (mean 50) possess compact ovaries
containing ovocytes of approximately the same size. The oviducts
are firm, flat and without convolution (fig. 1, open squares).

■ • • •

■ * • •

• • ■ •

• • • i

SNOUT-VENT

Figure 1. Ovarian condition correlated with body length. Each
square represents one individual. Open squares indicate ovaries
compact, oviducts straight. Squares with diagonals, ovaries
contain luteal-like structures and oviducts convoluted. Dotted
squares indicate compact ovaries and convoluted oviducts, and
represent specimens taken in August. All other individuals
were collected in July.

Twenty-three individuals in the size range 53 to 61 mm (average
56) possess one feature in common, the oviduct is always convoluted
(at least one loop is present) and, in the majority of individuals, this
convolution occurs in the proximal end. The ovaries of the July
1950 individuals (squares with diagonals, fig. 1) vary in the number
of collapsed follicles present. These same structures have been re-
ferred to as "corpora lutea" by McCurdy (1931), and presumably
are the remains of former ovarian follicles after ovulation has occurred
(McCurdy, op. cit.; Fisher and Richards 1950). Three individuals
with snout- vent lengths of 54, 58 and 56 mm possess no collapsed
follicles. The ovocytes are of two distinct sizes and the oviducts are
convoluted. All of the females collected in August 1952 (dotted
squares, fig. 1) possess ovaries which have completely recovered, ijs.
no collapsed follicles are present, ovocytes are of two distinct sizes,

No. 4 Gordon: A Population of Holbrook's Salamander 57

but the oviducts are convoluted.

The possession of a convoluted oviduct, at least prior to the egg-
laying season, seems to indicate that the individual has ovulated at
least once. This feature, plus the varying degrees of recovery from
ovulation, permits us to divide the sample into two groups, those in-
dividuals which have ovulated and presumably are sexually mature,
and a group which is becoming sexually mature and will be ready
for ovulation during the next reproductive season.

This interpretation is in sharp contrast with that made by Pope and
Pope (1949) for Pletbodon glutinosus glutinosus. They report a
group of twelve "spent" individuals (collected in late June, late July
and the first week in August) having ovaries which were not com-
pact, the eggs being of various sizes, and whose oviducts were not
swollen or convoluted. The body lengths of these individuals are
shorter than (yet overlap those of) a series of eleven obviously gravid
specimens (Pope and Pope, op. cit., fig. 63). The reproductive sea-
son is extended, presumably from June, or late May, through Septem-
ber (Pope and Pope, op. cit.).

The testis and vas deferens do not lend themselves as nicely to
macroscopic examination as do the female structures. Francis (1934)
reports that the Mullerian and Wolffian ducts are never pigmented
in female Salamandra while ". . . Muller's duct, and its associated
urino-genital duct . . ." are always pigmented in the male. Goin
(1950) states that the sex of Amby stoma c. bishopi may be deter-
mined ". . . by examination of the Mullerian duct, which is pig-
mented in the males and unpigmented in the females . . .". Pig-
mentation appears macroscopically to be confined to that portion of
the vas deferens (urino-genital or Wolffian duct) from which Muller's
duct is indistinguishable macroscopically (Francis, op. cit.) in mature
male Eurycea. Twenty-five males in the size range 43 to 54 mm
(average 50) possess vasa deferentia of which less than 50 percent
of the surface area is pigmented (see fig. 2). Of these, eight individ-
uals (average 50, extremes 48 to 54 mm) have no pigmentation, or
melanophores are faintly noticeable bilaterally along a line near the
dorsal surface of the duct. The duct in all of the twenty-five males
is firm and straight, exhibiting no convolutions. Sixteen specimens,
ranging from 51 to 58 mm, mean 54, have vasa deferentia which
are convoluted but vary from complete to partial pigmentation, al-
ways exceeding 50 percent of the surface of the duct. In this latter
group, pigmentation is absent from the proximal part of the duct, the
distal part being pigmented and convoluted. From the study of all
individuals, the spread of pigmentation seems to be from the distal
to the proximal end of the duct.

A general increase of pigmentation with size is suggested; this
may possibly be correlated with age and sexual maturity. If it is
correlated with the latter, then, as in the females, we have two groups
of animals, one (diagonal squares, fig. 2) sexually mature, having
been sexually active in one reproductive season; the other group (open
squares, fig. 2) just becoming sexually mature, will be ready for
sexual activity during the next reproductive season.

Sexual maturity is apparently reached at the same size in both sexes.

Tulane Studies in Zoology

Vol. 1

/ \

•»5

SNOUT-VENT

Figure 2. Condition of vasa deferentia correlated with body length.
Open squares indicate vas deferens pigmented less than 50
percent, no convolutions. Squares with diagonals, vas deferens
pigmented 50 percent or more and convoluted. Each square
represents one individual.

All evidence in the literature points to egg deposition in December
since females collected in November contain large ovarian eggs
(Brimley 1896, 1939; Parker 1948). Females collected in August
1952 possess ovocytes of two distinct sizes and represent the largest
individuals in the August sample. The largest females collected in
the July 1950 sample show varying degrees of recovery from the
"spent" ovarian condition. It would appear that egg deposition occurs
in late autumn for this Florida population, possibly in December. It
is interesting to note the length of the recovery phase from ovulation
until maturing ovocytes again make up the bulk of the ovary.

Vomerine Teeth
Counts were made on 144 individuals. Since only 39 percent of
the specimens have the same number of teeth on both sides, the sum
of the right and left counts is used in all calculations (see Pope and
Pope, 1949). Two individuals were discarded since the vomerine
teeth were not clearly separable from those of the parasphenoid.
There is no significant difference between the sexes in the number
of vomerine teeth. An increase in number of teeth with size is in-
dicated. Males less than 40 mm (snout-vent) have an average of
14.4 teeth, while females of a similar size group possess averagely
14.2 teeth. Of the individuals measuring more than 40 mm, males
average 19-8 teeth and females 19-7 teeth.

Body Measurements
The means and extremes for the various body measurements are
presented in Table 1. All measurements are in millimeters. Snout-
vent length is measured from tip of the snout to the anterior edge of
the vent. Head length is measured from the tip of the snout to the
gular fold. Head width is a measurement taken across the head at
the angle of the jaws. The axilla-groin measurement is the distance
between the limb insertions.

No. 4 Gordon: A Population of Holbrook's Salamander 59

Table 1. Body measurements.
No. Snout-vent Axilla-groin Head length Head width

Sexually mature

51.4 28.1

11.5

8.5

43-58 23-32

7-13

7-9

52.9 29.6

11.6

8.0

46-61 24-35

10-14

7-9

Sexually immature

34.2 18.3

8.3

5.7

29-37 16-20

7-9

5-6

34.6 18.6

8.5

5.8

31-39 16-21

8-9

5-7

The mature female reaches a total length only slightly larger than
the male. The average length for thirty-nine females is 148 mm
(extremes 122 to 167 mm). Thirty-four mature males average 147
mm in total length (extremes 125 to 165 mm).

Population Structure
Sex ratio. — The sex ratios do not deviate significantly from an ex-
pected 1 to 1 ratio of either the 1950 sample taken as a unit, the
1952 sample, or the two distinct size groups represented by both
samples combined.

Table 2. Sex ratios.

Sex 1950 1952 Juvenile Adult

Size groups. — The July 1950 sample (open squares, fig. 3) is com-
posed (with a single exception) of individuals in the size group 43
to 58 mm (snout-vent). These specimens as indicated above are
either sexually mature or would presumably have been mature by the
December 1950 reproductive season.

Parker (1948) reports that larvae collected in April measure 31
to 34 mm total length. Bishop (1943) gives the measurements of
larvae collected June 18 as varying from 22 to 39 mm. The measure-
ments of two recently transformed individuals are reported by Sinclair
(1951) as 22.5 and 24 mm body length, and 41.5 and 45 mm total
length. The single juvenile in the July 1950 sample measures 21
and 45 mm for the two lengths, and probably represents a recently
metamorphosed individual.

The body length frequency of the August 1952 sample presents a
distinctly different picture (squares with diagonals, fig. 3). Only
eight individuals are sexually mature. Fifty-five specimens (average
34, extremes 29 to 39 mm) are sexually immature and apparently
represent individuals metamorphosing the previous June or July.

If we assume that sexual maturity is reached in the second repro-
ductive season following metamorphosis, i.e. when the salamander is
two years old, then the unusual distribution of size groups in the two
samples may be explained in the following way.

It may be postulated that the collection of a large segment of
adults in July 1950, by reducing egg deposition the following De-
cember, would reduce the number of sexually maturing individuals
In 1952. This would explain the absence of the size group 42 to 55

Tulane Studies in Zoology

Vol. 1

□.

21.

■29

SNOUT-VENT

Figure 3. Frequency distribution — snout-vent length. Open squares
indicate specimens collected July 1950. Squares with diag-
onals indicate specimens collected August 1952. Each square
represents one individual.

mm in August 1952. The adults which reproduce in 1951 are
products of the eggs of 1949. Since the 1950 collection did not
interfere with deposition of eggs in 1949, nor with the newly meta-
morphosed individuals from these eggs, adults would be present to
reproduce in December 1951. Egg deposition occurred and the
larvae transformed in June or July. The August 1952 sample reveals
a preponderance of this group of metamorphosed individuals.

No obvious explanation is available for the absence of a larger
segment of the adults which bred in December 1951 from the August
1952 sample; however, differential habitat selection is known to
occur in many terrestrial salamanders and may be a possible explana-
tion in this instance.

References Cited

Bishop, Sherman C. 1943. Handbook of Salamanders. Comstock
Publishing Company, Ithaca, pp. 1-555.

Brimley, C. S. 1896. Batrachia found at Raleigh, N. C. Amer. Nat.,
30: 500-501.

1939. The amphibians and reptiles of North Caro-
lina. Carolina Tips [Elon College, N. C], 2: 18.

Carr, Archie F., Jr. 1940. A contribution to the herpetology of
Florida. Univ. Fla. Stud., Biol. Sci. Ser., 3: 1-118.

Chaney, Allan H. and Clarence L. Smith. 1950. Method for col-
lecting mapturtles. Copeia, 1950: 323-324.

Fisher, Helen T. and A. Richards. 1950. The annual ovarian
cycle of Acris crepitans Baird. In Researches on the Amphibia
of Oklahoma, Univ. Okla. Press, Norman, pp. 129-142.

Francis, Eric B. T. 1934. The Anatomy of the Salamander. The
Claredon Press, Oxford, pp. 1-381.

Goin, Coleman J. 1950. A study of the salamander, Ambystoma
cingulatum., with the description of a new subspecies. Ann.
Carg. Mus., 31: 299-320.

McCurdy, Harriet M. 1931. Development of the sex organs in
Triturus torosus. Amer. Jour. Anat., 47: 367-403.

Parker, Malcolm V. 1948. A contribution to the herpetology of
western Tennessee. Jour. Tenn. Acad. Sci., 12: 20-30.

Pope, Clifford H. and Sarah H. Pope. 1949. Notes on the growth
and reproduction of the slimy salamander, Plethodon glutinosus.
Fieldiana, 31: 251-261.

Sinclair, Ralph M. 1951. Notes on recently^ transformed larvae of
the salamander Eurycea longicauda guttolineata. Herpetol., 7 : 68.

Tinkle, Donald W. 1952. Notes on the salamander, Eurycea longi-
cauda guttolineata, in Florida. Field and Lab., 20 : 105-108.

HF\-\H^eu> Or\ts»

Volume 1, Number 5

September 1, 1953

A REDESCRIPTION OF THE CRAWFISH PROCAMBARUS

HINEI (ORTMANN)
(Decapoda, Astacidae).

GEORGE HENRY PENN,

DEPARTMENT OF ZOOLOGY, TULANE UNIVERSITY
OF LOUISIANA, NEW ORLEANS.

Mos. ctrap. mi

LIBRARY

SEP 4 195:3

HARVARD
UNIVERSITY

TULANE UNIVERSITY
NEW ORLEANS

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Horton H. Hobbs, Jr., Associate Professor of Biology, Univer-

of Virginia, Charlottesville, Virginia.
Fenner A. Chace, Jr,, Curator, Division of Marine Invertebrates,

United States National Museum, Washington, D. C.
Waldo L. Schmitt, Head Curator, Department of Zoology,

United States National Museum, Washington, D. C.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
torial matters, and orders for individual numbers to the editor. Re-
mittances should be made payable to Tulane University.

When citing this series authors are requested to use the following
abbreviations: Tulane Stud. Zool.
Price for this number: $0.25.

George Henry Penn, Editor,
c/o Department of Zoology,
Tulane University of Louisiana,
New Orleans, U. S. A.

ON OF THE CRAWFISH PROCAMBARUSl ,?£p-
HINEI (ORTMANN)

(Decapoda, Astacidae).

GEORGE HENRY PENN, pEP 4 19

Department of Zoology, Tulane University of Louisiana, "flRVflftD

New Orleans. 1/NlVfRSIT]

The crawfish which is the subject of this paper was named in honor
of the late James Stewart Hine (1866-1930), a distinguished student
of the Diptera, who was a summer visitor at the now non-existent
Gulf Biologic Station, Cameron, Louisiana. Dr. Hine sent the type
specimens to Dr. Arnold E. Ortmann at the Carnegie Museum who
described the species in 1905. Until recently this species was known
only from Cameron as recorded by Ortmann (1905), Cary (1906)
and Cary and Spaulding (1909); the latter two references are merely
repetitions of Ortmann's original data. In 1939 under a grant from
the New Orleans Academy of Sciences I collected crawfishes in
southern Louisiana and found this species in several additional parishes.
Later, Hobbs (1945) recorded it from Liberty County, Texas as a
co-inhabitant at the type locality of Cambarellus puer Hobbs. The
distribution and ecology of Procambarus hinei will be discussed at
length in a subsequent paper on the Louisiana species of the genus
Procambarus.

Ortmann's original description was inadequate and the figure of
the first pleopod of the form I male left much to be desired. Speci-
mens used in drawing up the following redescription were collected
in two areas in Cameron Parish, Louisiana, each within fifteen miles
of the type locality.

PROCAMBARUS HINEI (ORTMANN)

Synonymy. — Cambarus (Cambarus) hinei Ortmann, 1905, Ohio
Nat., 6: 401; Procambarus hinei (Ortmann), Hobbs, 1942, Amer.
Midi. Nat., 28: 342.

Type locality. — "One quarter mile from Gulf Beach, near Cameron,
Cameron Parish, Louisiana"; type unknown (or lost). The types are
not in the Carnegie Museum along with the rest of Ortmann's material
(Brooks, 1931), nor have they been located in the crawfish collections
of any other institution.

Male, form I. — Cephalothorax (figs. 1, 2) subovate. Abdomen
narrower than and slightly longer than the cephalothorax. Width of
cephalothorax at widest point subequal to height at the same point.

Areola broad, about 3!/2 times longer than width at its narrowest
point (average, 3.47; range, 3.78 to 2.86), with about five very fine
punctations in narrowest part (average, 5.5; range, 5 to 7). Cephalic
portion of cephalothorax about 2V4 times as long as the areola (aver-
age, 2.26; range 2.15 to 2.44); length of areola averages 30.6 percent
of the total length of the cephalothorax (range, 29-0 to 31.7 percent).

64 Tulane Studies in Zoology Vol. 1

Rostrum without lateral spines, but with margins interrupted in
most specimens; less than ten percent of the specimens with very
small lateral spines on rostrum. Rostrum widest at base, margins
raised, more or less straight and converging; no median carina. Acu-
men distinct, although its base merges with the remainder of the
rostrum.

Postorbital ridges weakly developed, terminating anteriorly in small
spines. Branchiostegal spine small, acute. Cervical groove inter-
rupted by a very small lateral spine on each side. In some specimens
this spine is absent. Epistome (fig. 3) broader than long, with a
small spine on anterior margin.

Cephalic region of the telson with two spines in each caudolateral
angle, the more lateral one nearly twice the length of the other.

Antennules of usual form, with a large spine on the distal margin
of the ventral surface of the basal segment. Antenna reaching just
beyond the telson. Antennal scale (fig. 4) extending well beyond
the tip of the rostrum; lateral margin straight, terminating in an
acute spine; lamellar portion flat, fringed with long hairs from base
to apex; greatest width of scale just proximal to the middle, length
about 2J4 times greatest width (average, 2.23; range, 2.00 to 2.50).

Chela (fig. 5) subcylindrical, long and slender; non-tuberculate;
very finely pubescent; palm inflated. Both fingers terminating in
short corneous tips bent toward each other. Opposable margins of
fingers flattened and covered with about four (apical) to about ten
(basal) rows of small rounded tubercles. Fingers very short; dactyl
about 35 percent of the total length of the outer margin of the chela
(average, 35.2; range, 32.0 to 37.0). Carpus (fig. 5) subcylindrical,
non-tuberculate.

Hooks (figs. 6,1) on ischiopodite of third and fourth pereiopods;
those of the fourth about two-thirds as long as those on third
pereiopods.

Venter of cephalothorax covered with a dense mat of long hairs
which are present on the mesial faces of all of the pereiopods and
maxillipeds. They are especially thick and conspicuous on the mesial
faces of the third maxillipeds and the first three pereiopods.

First pleopod (figs. 8, 9, 10) extending to caudal side of the
coxopodite of the third pereiopods when the abdomen is flexed.
Apical third of pleopod bent caudad at about a 60° angle to the
shaft. Pleopod terminating in three very small parts. The apical
elements are somewhat twisted counterclockwise so that the mesial
and cephalic processes are shifted to a more posterior position than
is normal; hence, they are most clearly seen when the caudal surface
of the pleopod is examined. Mesial process non-corneous, spiculi-
form and extending laterodistad; cephalic process corneous, lying
near to and mesiocaudad of the mesial process, truncate near apex,
and extending laterodistad; central projection corneous, acute and
somewhat compressed along its longitudinal axis, with fusion line of

No. 5

Penn: Redescription of Proca?nbarus hinei

Figures 1-15. Procambarus hinei (Ortmann) : 1, 2, cephalothorax
of form I male; 3, epistome of form I male; 4, antennal scale of
form I male; 5, chela and carpus of form I male; 6, 7, hooks on
ischiopodites of third and fourth pereiopods of form I male; 8, 9,
10, mesial, caudal and lateral views of the first pleopod of form I
male; 11, 12, 13, mesial, caudal and lateral views of the first pleo-
pod of form II male; 14, chela and carpus of female; 15, annulus
ventralis of female. Pubescence removed from all structures illus-
trated.

66 Tulane Studies in Zoology Vol. 1

its component elements clearly visible. Caudal process not developed.
A shoulder-like hump is present on the cephalomesial part of the apex
of the pleopod. Apical half of mesial surface of pleopod with a heavy
clothing of long hairs.

Male, form II. — Very similar to form I in general appearance.
Chelae reduced but with the ratio of length of fingers to length of
chela as in the form I male. Hooks on ischiopodites of third and
fourth pereiopods greatly reduced. Venter of cephalothorax only
sparsely clothed with long hairs. First pair of pleopods (figs. 11, 12,
13) reaching to posterior part of coxopodites of third pereiopods
when abdomen is flexed; apical processes reduced and non-corneous.

Female. — Similar to form I male in shape and structure of the
cephalothorax, but differing strikingly in the proportions of the chelae
(fig. 14) in which the palm is only slightly longer than the fingers.
Annulus ventralis (fig. 15) immovable, roughly pyramidal in shape
with the apex of the base directed anteriorly; width nearly twice as
great as length. Anterior face with a deep furrow from the base to
the summit of the pyramid along the anterior edge. Sinus originates
at the summit of this furrow and runs posterodextrad and gently curves
back posterosinistrad nearly to the base of the posterior face of the
pyramid.

Measurements. — Following are measurements in millimeters for the
largest and smallest of the form I males, and the largest female in the
collections examined.

(largest) £

)i (smallest

) $ (larg

Cephalothorax :

Length

20.0

14.5

22.0

Height (greatest)

9.9

7.1

11.0

Width (greatest)

9.8

7.6

11.0

Areola :

Length

6.0

4.3

7.5

Width (narrowest)

1.7

1.5

2.0

Rostrum:

Length

5.5

4.5

5.6

Width at base

3.1

2.2

4.2

Antennal scale:

Length (lateral margin)

5.2

4.3

5.5

Width (greatest)

2.5

1.9

2.7

Abdomen :

Length (to tip of tel'son)

27.0

18.5

27.0

Chela:

Length of outer margin

18.0

10.5

9.0

Length of inner margin

of palm

11.5

5.8

3.8

Width of palm (greatest)

4.5

2.4

3.5

Thickness of palm

(greatest)

3.5

2.0

2.4

Length of dactyl

5.9

3.9

4.5

Color pattern. — Although all of my specimens are faded so that the
exact color cannot be recognized, the following notes made some

No. 5 Penn: Redescription of Procambarus hinei 61

years ago will at least indicate the color pattern. Dorsally with a
pair of conspicuous dark, broad, parallel, longitudinal stripes which
originate just below the postorbital ridges and run posteriorly along
either side of the areola and onto the abdomen. On the abdomen
these stripes converge slightly, become narrower and terminate on
the base of the telson. At their termination the stripes are about half
as wide as at their points of origin. The sides of the abdominal
tergites each have a thin longitudinal, darker stripe. Chelae without
dark markings.

Specimens examined, — The specimens used in this study were col-
lected from two localities in Cameron Parish, Louisiana as follows:

5 $ Sj, 8 $ $ juveniles, 3 ??, and 7 9 9 juveniles, from a
shallow pond on the coastal chenier at Creole, April 22, 1940, P.
Viosca, Jr. and G. H. Penn (TU P-553). Included in the same col-
lection were a few each of Procambarus blandingii acutus (Girard)
and P. clarkii (Girard).

32 $ $ i, 9 $ $ n, and 14 S S juveniles, from a shallow pond at
Hackberry, July 28, 1940, G. H. Penn (TU P-557). Included in the
same collection were a few each of Cambarellus puer Hobbs and Pro-
cambarus clarkii (Girard).

Relationships. — Ortmann (1905: 403) placed P. hinei in the
Blandingii Section on the basis of the hooks on its third and fourth
pereiopods and the subcylindrical chelae, and in the Alleni Group be-
cause of the shape of its rostrum and width of its areola. He did
state, however, that "within the latter group it stands rather isolated
with regard to the male organs, which show a rather primitive con-
formation, with exception of the distinct backward curve of the
distal part." However, Hobbs (1942: 70-71) pointed out that P.
alleni (Faxon) itself is a very disjunct species, and he thus considers
the Alleni group to be monotypic and probably belonging in the
Barbatus Section. In view of the peculiarities of the structure of the
first pleopod of the form I male, Procambarus hinei cannot be placed
in the same group with P. alleni, nor even in the same section; neither
can it be placed logically in any of the other sections of the genus as
currently described. Accordingly, I am designating a separate section,
the Hinei Section, to accommodate this disjunct species.

REFERENCES CITED

Brooks, Stanley T. 1931. List of types of Crustacea in the collection
of the Carnegie Museum on January 1, 1931. Ann. Carnegie
Mus., 20: 161-167.

Cary, L. R. 1906. A contribution to the fauna of the coast of Louisi-
ana. Gulf Biologic Sta., Bull. No. 6 : 50-59.

Cary, L. R. and H. M. Spaulding 1909. Further contributions to the
marine fauna of the Louisiana coast. Gulf Biologic Sta., Bull.
No. 12: 3-21.

Hobbs, Horton H., Jr. 1942. The crayfishes of Florida. Univ. Fla.
Publ, Biol. Sci. Ser., 3(2) : 1-179.

68 Tulane Studies in Zoology Vol. 1

1945. Two new species of crayfishes of the genus

Cambarellus from the Gulf coastal states, with a key to the
species of the genus. Amer. Midi. Nat., 34: 466-474.

Ortmann, A. E. 1905. A new species of Cambarus from Louisiana.
Ohio Nat., 6: 401-403.

Penn, George H. 1941. Preliminary report of a survey of the craw-
fishes of Louisiana. [Abstract] New Orleans Acad. Sci., Abstr.
Pap. 88th Ann. Meet., 1941 : 8.

\ -

Volume 1, Number 6

September 15, 1953

A NEW BURROWING CRAWFISH OF THE GENUS

PROCAMBARUS FROM LOUISIANA AND

MISSISSIPPI

(Decapoda, Astacidae)

GEORGE HENRY PENN,

DEPARTMENT OF ZOOLOGY, TULANE UNIVERSITY
OF LOUISIANA, NEW ORLEANS.

TULANE UNIVERSITY
NEW ORLEANS

MUS. COMP. ZiOL

LIBRARY

SEP 1 8 1953
UNIVERSITY

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Horton H. Hobbs, Jr., Associate Professor of Biology, Univer-
sity of Virginia, Charlottesville, Virginia

Fenner A. Chace, Jr., Curator, Division of Marine Invertebrates,
United States National Museum, Washington, D. C.

Waldo L. Schmitt, Head Curator, Department of Zoology,
United States National Museum, Washington, D C.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
torial matters, and orders for individual numbers to the editor. Re-
mittances should be made payable to Tulane University.

When citing this series authors are requested to use the following
abbreviations: Tulane Stud. Zool.

Price for this number: $0.25.

George Henry Penn, Editor,
c/o Department of Zoology,
Tulane University of Louisiana,
New Orleans, U. S. A.

LIBRARY

SEP 1 8 195

HARVARD
UNIVERSITY

A NEW BURROWING CRAWFISH OF THE GENUS

PROCAMBARUS FROM LOUISIANA AND

MISSISSIPPI

(Decapoda, Astacidae)

GEORGE HENRY PENN,

Department of Zoology, Tulane University of Louisiana,

New Orleans.

The new species of Procambarus described herein occurs over a
fairly wide geographic area, however, it has never been found in large
numbers and is a rarely encountered form. Ecologically it may be
included among the secondary burrowers (Hobbs, 1942: 20). The
relationships of this new species are not clear and will be discussed
following the description.

PROCAMBARUS PLANIROSTRIS, sp. nov.

Holotype male, form I. — Body subovate, appearing somewhat com-
pressed; abdomen about equal to length of cephalothorax ( 29.0 - 27.5
mm). Height of cephalothorax (figs. 1, 2) slightly less than width
in region of caudodorsal margin of the cervical groove (12.5-13.0
mm); greatest width of cephalothorax slightly caudad of caudodorsal
margin of the cervical groove.

Areola narrow (20 times longer than width), with a single punc-
tation in the narrowest part; cephalic portion of cephalothorax about
1.75 times as long as the areola; length of areola about 36 percent of
total length of cephalothorax.

Rostrum without lateral spines; widest at base, margins slightly
raised and only slightly thickened, converging at acumen. Upper
surface almost flat, moderately punctate. Acumen small, directed
dorsally at tip.

Postorbital ridges reduced, terminating anteriorly without spines;
lateral surface excavate. Branchiostegal spine small. Cervical groove
interrupted laterally; lateral spine reduced to the size of a large
tubercle. Lateral surfaces of cephalothorax granulate, dorsal surface
moderately granulate.

Cephalic region of telson with spines in each caudolateral angle,
three on right, five on left.

Epistome (fig. 3) slightly more than twice as wide as long, with
slightly concave center; cephalic margin with a small spine.

Antennae nearly equal to total length of the crawfish; of normal
form. Antennal scale (fig. 4) narrow; widest a little distad of middle;
lateral margin inflated, straight and terminating distally in a small
spine; total length less than length of areola (8.5 - 10.0 mm).

Chela (fig. 5) with palm inflated; fingers slightly depressed; seti-
ferous punctations present over dorsal surface of most of palm and
both fingers. Inner margin of palm with a row of eight prominent

72 Tulane Studies in Zoology Vol. 1

tubercles. Both fingers terminating in short corneous tips, that of
the dactyl overhanging the other when the fingers are closed. Thir-
teen rounded tubercles at base and one distally-located corneous tuber-
cle on opposable margin of immovable finger; fourteen rounded
tubercles in corresponding positions on the dactyl. Upper surface of
dactyl with seven strong tubercles basally.

Carpus (fig. 5) with five strong spines on distal end in a semi-
circular arrangement extending medioventrally from dorsal to ventral
condyles which articulate with the chela. Upper surface with smaller
tubercles toward inner margin and scattered setiferous punctations
generally.

Simple hooks (fig 6) present on ischiopodites of third and fourth
pereiopods; length of each greater than half the diameter of the
respective ischiopodites.

First pleopod ( figs. 7, 8, 9 ) reaching to anterior side of the coxopo-
dite of the third pereiopods when the abdomen is flexed. Apex
terminating in four distinct parts which as a unit extend cuadad at
about a 40° angle to the shaft of the pleopod. Mesial process non-
corneous, spiniform, directed caudodistad, and not extending beyond
the other terminal parts; cephalic process corneous, arising on mesial
side of the central projection, directed caudodistad, excavate on caudo-
lateral surface, and closely applied to the central projection; central
projection corneous, compressed, "beak-like" in shape, with apex
directed caudad; fusion line of centrocaudal and centrocephalic com-
ponents clearly indicated. Caudal process consisting of two corneous
parts: mesially a low, longitudinal ridge flanked laterally by a leaf-
like element which extends distally, its apex coming in contact with
the overhanging central projection. Cephalic margin of the shaft of
the pleopod with a distinct shoulder.

Morpbotype male, form II. — Very similar to holotype in general
appearance; chelae and hooks on ischiopodite of the third and fourth
pereiopods greatly reduced. First pair of pleopods (figs. 10, 11)
reaching to middle of coxopodites of third pereiopods when the ab-
domen is flexed; all processes reduced and non-corneous.

Allotype female. — Very similar to holotype in general appearance;
chelae greatly reduced. Annulus ventralis (fig. 12) immovable,
roughly spindle-shaped with center produced into a cone-shaped pro-
tuberance on either side of which is a shallow groove. The sinus
originates on the center line a short distance removed from the an-
terior margin then proceeds through a gently zigzag course to the
apex of the central, cone-shaped protuberance. The sternum of the
preceding thoracic segment is slightly produced so that its posterior
margin underhangs the anterior margin of the annulus.

Color. — The following color notes were made from living mature
specimens collected at the type locality. In general this is a drab-
colored species. The effect is that of a light tan overcast with olive

No. 6

Venn: New Burrowing Crawfish

Figures 1-12. Procambarus planirostris, sp. nov. : 1, 2, cephalo-
thorax of the holotype; 3, epistome of the hclotype; 4, antennal
scale of the holotype; 5, chela and carpus of the holotype; 6, hooks
on ischiopodites of the third and fourth pereiopods of the holotype;
7, 8, 9, mesial, caudal and lateral views of the first pleopod of the
holotype; 10, 11, mesial and lateral views of the morphotype; 12,
annulus ventralis of the allotype. Pubescence removed from all
structures illustrated.

74 Tulane Studies in Zoology Vol. 1

dorsally on the cephalothorax; the rostrum tan only. Abdomen
dorsally with an inconspicuous wide stripe of olive-tan which tapers
to a point on the base of the telson; background color of abdomen on
either side of the dorsal stripe is a light tan with very fine flecks of
darker reddish-tan; this background color extends also onto the dorsal
surface of the telson and uropods. On the dorsolateral parts of the
abdominal tergites there is a row of spots of olive, one on the anterior
margin of each tergite, and connected longitudinally by a faint line
of olive. Chelae of the same basic color as the background color of
the abdomen, but tubercles capped with black or dark brown, giving
the chela and carpus a fine-spotted appearance. One specimen had
a faint bluish cast to the fingers.

Measurements. — As follow:

;, in milhmt

:ters:

Holotype

Allotype

Morphotype

Cephalothorax :

Length

27.5

20.0

29.0

Width (greatest)

13.5

10.0

14.0

Height (greatest)

12.5

9.5

13.5

Areola :

Length

10.0

7.5

10.5

Width (at narrowest point)

0.5

0.7

Rostrum :

Length

5.5

4.0

6.5

Width at base

5.5

3.5

5.5

Abdomen :

Length (to tip of telson)

29.0

22.0

29.0

Right chela :

Length of outer margin

of hand

25.5

11.0

21.0

Length of dactyl

15.0

6.0

12.0

Width of palm (greatest)

8.5

4.0

7.5

Thickness of palm

(greatest)

6.0

2.5

5.0

Length of inner margin

of palm

10.0

4.0

8.0

Type locality. — The holotype and allotype were collected from a
low area of mixed hardwood, pine and palmetto flatwoods one mile
south of Walker (on Louisiana highway 336), Livingston Parish,
Louisiana. The holotype was taken on February 17, 1951 by Dr.
R. D. Suttkus when the area was inundated by about a foot of water;
the allotype was collected on August 29, 1952 by the author and
C. E. Biggs from a simple burrow with a neat chimney, around the
base of which there was about six inches of standing water. The
soil here is a whitish clay and the burrow extended to about twelve
inches beneath the soil surface. At the same place there were
numerous burrows of Cambarus hedgpethi Hobbs and Orconectes
clypeatus (Hay).

The morphotype was collected from a small creek three miles south
of Janice, Perry County, Mississippi on January 28, 1951 by Dr. Fred
R. Cagle. No other crawfishes were found at this locality.

No. 6 Venn: New Burrowing Crawfish 75

Disposition of types. — The holotype, allotype and morphotype are
deposited in the United States National Museum, catalogue numbers
95674, 95675, and 95676 respectively. The paratypes are in the fol-
lowing collections: Academy of Natural Sciences, Philadelphia (1 $ If
2 $ $ juv., and 1 ? ) , the personal collection of Dr. Horton H.
Hobbs Jr. at the University of Virginia (1 $ b 1 $ juv., and 1 $ ) ,
and Tulane University (2 ^,,7 t $ juv., 4 2 9, and 6 5 9 juv.).

Geographic distribution. — The type series of Procambarus plani-
rostris was collected from the "Florida" parishes of southeastern
Louisiana and southern Mississippi. These records and a summary
of the deposition of these specimens are as follows. LOUISIANA:
East Baton Rouge Parish: 9 mi. s. Baton Rouge, January 26, 1949,
G. H. Bick and L. L Ellis (TU 910); Livingston Parish: 1 mi. s.
Walker, February 17, 1951, R. D. Suttkus (USNM, TU 2278), same
locality, July 19, 1952, G. H. Penn, R. D. Suttkus and C E. Biggs
(ANS, HHH), same locality, August 20, 1952, G. H. Penn and
C. E. Biggs (USNM); St. Tammany Parish: Lake Pontchartrain at
Mandeville, February 22, 1935, P. Viosca, Jr. and H. B. Chase (TU
P-610); Washington Parish: 6 mi. nw. Enon, August 10, 1948,
G. H. Penn and M. H. Penn (ANS), Franklinton, March 27, 1949,
F. R. Cagle (ANS, HHH), 2 mi. n. Varnado, March 3, 1953, F. R.
Cagle (TU 2894). MISSISSIPPI: Perry County: 3 mi. s. Janice,
January 28, 1951, F. R. Cagle (USNM, TU 2853).

Relationships. — Procambarus planirostris appears not to belong defi-
nitely in any of the sections of Procambarus as currently recognized,
but has certain characteristics of each of two sections, and apparently
occupies a somewhat intermediate position between the two. It shows
affinities with the Barbatus Section (Hobbs, 1942: 35-36) in its gen-
eral body conformation and in that the cephalic process of the first
pleopod arises from the mesial side of the central projection, but
differs from members of this section in lacking the accessory cephalo-
distal ridge or knob-like prominence. It shows closer affinities with
the Clarkii Subgroup of the Blandingii Section as defined by Hobbs
(1942: 93, 98-99) in possessing a distinct shoulder on the cephalic
margin of the first pleopod, and in the general configuration of the
annulus ventralis.

Because P. planirostris exhibits this peculiar combination of char-
acteristics a new section to accommodate it could be justified. How-
ever, because of its assumed closer relationship to the species of the
Clarkii Subgroup of the Blandingii Section than to those of the Bar-
batus Section, I am placing it in a separate subgroup, the Planirostris
Subgroup in the Blandingii Section. This action necessitates the modi-
fication of Hobbs' diagnosis (1942: 93) with regard to the cephalic
process of the first pleopod to the following: cephalic process when
present arises from cephalic or cephalolateral margin in all species
except those of the Planirostris Subgroup in which it arises from the
mesial side of the central projection.

76 Tulane Studies in Zoology Vol. 1

REFERENCE CITED

Hobbs, Horton H., Jr. 1942. The crayfishes of Florida. Univ. Fla.
Publ, Biol. Sci. Ser., 3(2) : 1-179.

V 19 IDS 3

Volume 1, Number 7

October 23, 1953

THE LIFE HISTORY OF THE CRAWFISH

ORCONECTES (FAXONELLA) CLYPEATUS (HAY)

(Decapoda, Astacidae)

ELSIE WAYNE SMITH,

DEPARTMENT OF ZOOLOGY. TCLANE UNIVERSITY,
NEW ORLEANS.

VMS. COM?. lmi~

OCT 2 7 1953
MUM

mmmw

TULANE UNIVERSITY
NEW ORLEANS

TULANE STUDIES IN ZOOLOGY is devoted primarily to the
zoology of the area bordering the Gulf of Mexico and the Caribbean
Sea. Each number is issued separately and deals with an individual
study. As volumes are completed, title pages and tables of contents
are distributed to institutions exchanging the entire series.

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Horton H. Hobbs, Jr., Associate Professor of Biology, Univer-
sity of Virginia, Charlottesville, Virginia.

Thomas Park, Professor of Zoology, University of Chicago,
Chicago, Illinois.

GEORGE H. Penn, Associate Professor of Zoology, Tulane Uni-
versity, New Orleans, Louisiana.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
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mittances should be made payable to Tulane University.

When citing this series authors are requested to use the following
abbreviations: Tulane Stud, Zool.

Price for this number: $0.50.

George Henry Penn, Editor,

c/o Department of Zoology,
Tulane University,
New Orleans, U. S. A.

THE LIFE HISTORY OF THE CRAWFISH
ORCONECTES (FAXONELLA) CLYPEATUS (HAY)

(Decapoda, Astacidae)

ELSIE WAYNE SMITH,i

Department of Zoology, Tulane University,

New Orleans.

The details of the life histories of comparatively few species of
crawfishes in the United States have been studied, however the life
history of at least one species in each of the four major genera is fairly
well known. Within the genus Orconectes two species of the subgenus
Orconectes have been well studied: 0S propinquus propinquus (Gi-
rard) by Creaser (1933a) in Michigan, and by Van Deventer (1937)
and Bovbjerg (1952) in Illinois; and O. immunis immunis (Hagen)
by Tack (1941) in New York. The subject of the present study, 0.
clypeatus, a member of the subgenus Faxonella, has a somewhat dis-
tinctive life cycle which might prove to be characteristic of this sub-
genus alone.

When this study was begun practically nothing had been published
concerning either the ecology or life cycle of the species. The entire
literature may be summarized in a few sentences. Hay (1899) de-
scribed it as Cambarus clypeatus on the basis of a single female col-
lected from a skiff at Bay St. Louis, Hancock County, Mississippi.
From the date of this inauspicious first scientific recognition, cly-
peatus remained a species of questionable affinities (c.f. Faxon, 1914)
until Creaser (1933b) described the first and second-form males and
redescribed the female from two series of specimens collected in
Louisiana and Alabama. He also included brief notes describing the
"pools along the roadside" and "pools in a bog or swamp" respectively
in which it was collected and included the statement that "many bur-
rows were found along the edge of these pools ... no doubt this
species is a burrower, for the pools certainly dry at certain seasons of
the year." Later the same year, Creaser and Ortenburger (1933)
recorded O. clypeatus from Oklahoma and repeated Creaser's notes on
its ecology. Lyle (1938) recorded it from Mississippi without com-
ment. Hobbs (1942) recorded it from Florida, Georgia, Alabama
and Arkansas and described its burrows along the sides of a roadside
ditch in Jackson County, Florida as "marked by small, neatly con-
structed chimneys, simple, and ranging in depth from six inches to a
foot." Penn (1942) recorded the species as abundant in a "pine
barren pond" in Louisiana. Finally, Penn (1952) published a com-
pilation of the data of the distribution and ecology of the species in
Louisiana.

Within the boundaries of Louisiana O. clypeatus is one of the com-
monest species of crawfishes, and occurs in the following variety of
habitats as listed by Penn (1952); temporary situations (58%), in-
cluding roadside ditches, pineland sloughs, puddles and borrow pits,
potholes in dry creek beds, and burrows; permanent situations (42%),

1 Present address: Savannah, Georgia.

MUS. C
LIBRARY

OCT 2 7 19!

HARVARD

UNIVERSITY

80 Tulane Studies in Zoology Vol. 1

including ponds, creeks, rivers, swamps and swamp ponds. Within
the state it occupies these habitats in all physiographic regions except
the recent alluvial lands and coastal marshes. The overall distribution
of the species as presently known includes the Coastal Plain from
southeastern Oklahoma to northeastern Arkansas and Georgia.

THE STUDY AREA
The life history of 0. clypeatus was studied from October 1949
through January 1951 by means of field observations and statistical
analyses of periodic samples of a natural population. The population
which was studied and from which the samples were collected was
located in a pineland ditch 2.4 miles south of Hickory, St. Tammany
Parish, Louisiana. The ditch itself lay to the south side of an unused
logging road which formed a retaining levee for an extensive, perma-
nently inundated slough on its north side. The ditch was about 200
feet long, from one to two-and-one-half feet deep, and from three to
five feet wide. On the bottom and along the sides, the ditch was
covered with a thick carpet of Juncus repens Michx. throughout the
entire year, and at least some parts of the ditch were shaded through-
out each day.

Drought conditions, characterized by the complete absence of stand-
ing water anywhere in the ditch, and wet conditions characterized by
the presence of standing water throughout the ditch, both occurred
periodically during the study period (Table 1). During the wet
periods the water level fluctuated with the amount of local rainfall,
occasionally overflowing the banks. Usually either one or the other
situation obtained when sampling was done, but an intermediate con-
dition existed in which the deeper parts of the ditch were wet and
the shallower parts dry. At such times separate samples were col-
lected from each situation. Other crawfishes present in the ditch
were numerous Cambarellus shufeldtii (Faxon) and occasional Pro-
cambarus blandingii acutus (Girard).

METHODS OF STUDY
The primary approach to the study was the analysis of periodic
samples removed from the population. Collections of samples were
originally planned for four week intervals over a period of sixteen
months, but it was not practical to adhere strictly to this schedule. A
total of sixteen samples on fourteen dates was collected; however,
samples for six of the sixteen months are not represented (Table 1).
During November and December 1949, drought conditions prevailed
and, although field trips were made, the burrowing population was
not found. During the other four months, January, March, August
and November 1950, no field trips were made because of transporta-
tion difficulties.

The samples were collected by dipnet during the wet periods and
by digging during the dry periods. Both methods were used along
the sides of and in the middle of the ditch for each collection. From
one-fourth to one-third of the length of the ditch was sampled when
the ditch was wet, but during drought conditions the slower and more

No. 7

Smith: Life History of Orconectes clypeatus

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82 Tulane Studies in Zoology Vol. 1

laborious digging was restricted to from one-sixteenth to one-eighth
of the length of the ditch.

The first sample made from burrows was taken in the second dry
period (May 1950) after the study was begun. The burrows were
found over the bottom and along the sides of the ditch and each was
marked only by an inconspicuous entrance hole, usually less than
three-fourths of an inch in diameter; chimneys were never seen.
Below the entrance each burrow branched off into several tunnels,
which eventually led to individual pockets or "cells" from eight to
fifteen inches beneath the surface. Each pocket was occupied by one
crawfish only.

Although this collection established the fact that the population
burrowed during dry periods, the possibility that a portion of it might
migrate during such times remained to be determined. Accordingly,
each time a visit was made to Hickory, the slough across the road was
checked with a dipnet for a migrating part of the population. Only
six individuals of O. clypeatus were found in the slough and these
only during wet periods following times when the ditch could have
overflowed and flooded the road. If drought is a necessary factor for
the completion of the life cycle of this species then the facts that the
road was seldom flooded and that the unpopulated slough always con-
tained water explains the scarcity of this species in collections from
the slough. It was felt therefore that the absence of 0. clypeatus
from collections during the October-November 1949 drought v/as a
result of faulty technic in sampling, not of an absence of the burrow-
ing population.

The majority of the samples was preserved in alcohol either in the
field or immediately upon return to the laboratory. Some first-form
males, collected December 1, 1950, and some ovigerous females, col-
lected between September and December 1950, were kept alive for
observation in the laboratory.

In the laboratory the length of the cephalothorax was measured on
all specimens. This measurement was taken on an observed mid-
dorsal line from the tip of the rostrum to the end of the cephalothorax.
All measurements were made with an ocular micrometer calibrated
for a stereoscopic microscope. Each measurement was estimated to
the nearest part of a micrometer unit but was converted for recording
only to the nearest one-tenth (0.1) of a millimeter. As it was
measured and recorded each specimen was assigned a sub-lot number
and kept in an individual vial until the entire lot (i.e<, sample) had
been completely studied.

A series of females was selected for the study of ovarian egg growth
and production. After an ovary had been removed, the eggs were
teased from it in a dish of water. Eggs were measured along their
longest diameters and recorded to the nearest hundredth of a milli-
meter (0.01). Total ovarian egg count was recorded for only a few
of the females so analysed.

A few specimens, in addition to certain samples, were collected
separately and kept alive for observation in the laboratory. Such in-

No. 7 Smith: Life History of Orconectes clypeatus 83

dividuals were placed each in separate bowls or small aquaria con-
taining rain water, sand, and usually some vegetation. All were fed
Pablum.

STATISTICAL ANALYSES

A detailed discussion of the statistical analyses used in this study is
not included in this paper. However, since the conclusions and inter-
pretations of the life cycle are based largely on statistical methods, a
brief summary of the methods employed and the steps of the various
analyses is included here.

Methods used in the statistical analyses were modeled closely on
those given by Peatman (1947) and Cazier and Bacon (1949). The
measurement of cephalothorax length was taken as the criterion of
age, but sexual maturity was based on other well-established biological
criteria, *>., attainment of first form in the males, and maturation of
ovarian eggs or the ovigerous condition in the females.

Since it was anticipated that each sample would give a clue to the
stage of development of the entire population during a given time
fragment in its annual cycle, it was necessary first to determine the
homogeneity or extent of heterogeneity of the apparent age com-
ponents (i.e., size groups) of each sample. This was done following
the grouping of measurements according to the following criteria:
(1) each sample was kept separate according to time and place (i.e.,
either aquatic or burrowing); (2) females, first-form males, second-
form males, and unsexed young were separated into sub-groups re-
spectively whenever they occurred in any one sample. At this point
the sub-groups of the samples were each considered to be homogeneous
on the fundamentals of location, time, and sex, but not on age.

Determination of homogeneity of age of these groups was based on
three hypotheses: (1) that the frequency distribution of the cephalo-
thorax length describes the form of the normal curve when the group
is homogeneous; (2) that those groups which do not fit the normal
curve, do not because of the presence of more than one age group
within the group tested, and vice versa, those groups that do, do so
because of the presence of only one age group; and, (3) that the
heterogeneous groups (according to age only) can be resolved into
their homogeneous components under certain conditions.

Grouped data were used throughout the study for all calculations of
mean, standard deviation, and for those other statistics that were based
upon the mean or standard deviation. The criteria for class groupings
of the samples were: one-tenth millimeter (the very youngest craw-
fishes); two-tenths millimeter (the maturing groups), and five-tenths
millimeter (the adult groups).

The first hypothesis was proved by using a set of young crawfish
removed from the pleopods of their mother, thus of a known uniform
(or homogeneous) age (Table 2, Dec. 1, group "j"). The second
and third hypotheses, dependent on the proof of the first, were re-
solved by applying the chi -square Test of Significance to the separate
frequency distributions of each of the sub-groups already known to
be homogeneous except for age. The results (chi-square probabilities)

Tulane Studies in Zoology

Vol. 1

TABLE 2.

Basic Statistics for Groups of Females
and Unsexed Juveniles

Cephalothorax Lengtl

i in

Number of

Millimeters

Date

Group

Specimens -

Actual Range

Mean

1949:

Oct. 16

18.8

—

13.4 — 16.1

14.9

.50

1950:

Feb. 24

5.2— 5.8

333

6.0 — 10.0

8.0

.02

10.3 — 12.7

11.0

.50

13.8 — 18.6

15.7

.99

Apr. 9

7.8— 8.8

—

141

8.9 — 12.3

10.fi

.30

13.0 — 16.9

15.1

.50

15.5 — 18.2

16.7

.50

19.0 — 19.6

■ —

—

May 12

8.1

- —

—

9.8 — 13.3

11.5

.90

III

13.6 — 14.6

—

June 2

10.6 — 13.3

12.0

.90

June 13

8.6 — 10.2

148

10.5 — 13,8

12.2

.10

14.1 — 16.5

—

July 12

10.9 — 13.8

12.4

.50

July 15

9.6 — 14.4

12.4

.10

Sep. 29

12.3 — 15.4

13.7

,50

Oct. 15

4.8— 5.2

14.7

—

Oct. 27

3*»

2.1— 2.3

14.8

—

Dec. 1

50* *#

2.4— 2.8

2.6

.20

44**

2.0— 3.2

2.7

.50

o**

3.8— 4.2

—

11.2

—

13.1 — 15.2

—

Dec. 29

2.8— 3.4

148

3.5— 4.7

4.2

.50

11.5 — 18.0

14.2

.50

1951 :

Jan. 24

3.0— 3.4

3.2

.90

155

3.7— 4.6

4.1

.30

174

4.5— 5.8

5.2

.50

119

5.6— 6.6

6.1

.50

6.7— 7.0

10.8 — 11.7

. —

13.8 — 19.0

16.7

.02

* Mean and chi-square probability not calculated for groups of less than
15 specimens. A chi-square probability of 0.10 (10%) and above esti-
mates with confidence that the groups are homogeneous.

** Unsexed juveniles.

# Attached to pleopods of mother.

gave estimates only of the occurrence of significant deviations or the
lack of deviations of any of the distributions from the normal dis-
tribution. A significant deviation was interpreted as the result of the
inclusion of more than one age group within a distribution. Ac-
cordingly, the frequency distributions of the thus-established hetero-
geneous age groups were re-observed and subdivided into apparent
smaller homogeneous groups {i.e., at apparent modal breaks, or de-

No. 7

Smith: Life History of Orconectes clypeatus

TABLE 3.
Basic Statistics for Groups of Males

Group

Cephalothorax Length in

Millimeters

Number of -
Specimens

Date

Actual Range

Mean

1949:

Oct. 16

11.9 — 16.6

14.4

.50

13.7 — 13.9

19.2

- —

—

1950:

Feb. 24

3.7— 5.8

—

310

6.0 — 10.2

8.1

.50

10.4 — 14.8

12.4

.20

12.6 — 17.3

14.8

.09

Apr. 9

7.2— 8.5

168

8.9 — 12.4

10.4

.50

12.5 — 15.4

13.6

.50

16.1 — 16.4

—

11.9 — 14.8

13.6

.50

15.1 — 17.5

—

May 12

8.7

—

9.5 — 13.4

11.5

.50

III

14.2 — 14.7

—

11.5 — 13.8

—

June 2

9.8 — 13.8

11.6

.20

12.0 — 13.6

—

June 13

8.4— 9.8

153

10.0 — 13.2

11.7

.50

13.7 — 14.5

—

16.1

—

11.8 — 13.6

12.8

.20

July 12

11.1 — 13.1

12.1

.50

11.3 — 13.6

1 — ■

—

July 15

9.2

___

10.4 — 14.2

12.1

.50

15.3

—

11.0 — 14.2

12.5

.10

18.2

—

Sep. 29

12.8 — 14.6

10.8 — 15.3

12.9

.50

16.4

—

Oct. 15

13.0 — 13.8

—

Oct. 27

14.0

11.9 — 14.1

—

16.4

—

Dec. 1

12.9 — 13.3

13.5

—

Dec. 29

3.3— 3.6

140

3.7— 4.6

4.2

.50

10.7 — 15.8

13.3

.50

11.8 — 14.5

—

^f^K'1 •

16.6

—

1951 :
Jan. 24

3.0— 3.6

3.4

.90

146

3.6— 4.4

4.0

.50

4.5— 5.1

5.0

.95

121

5.2— 6.1

5.7

.30

6.1— 6.8

6.4

.50

13.4 — 13.9

—

14.0 — 18.4

15.8

.90

11.6 — 18.5

14.2

.50

* Mean and chi-square probability not calculated for groups of less than
15 specimens. A chi-square probability of 0.10 (10%) and above esti-
mates with confidence that the groups are homogeneous.

86 Tulane Studies in Zoology Vol. 1

pressions in the frequency distributions) and the chi-square test
applied to each new sub-group. In the majority of cases the hetero-
geneous age groups could be subdivided by this procedure into homo-
geneous age groups with an acceptable degree of confidence.

When all lots had been subdivided into their component homo-
geneous age groups (Tables 2 and 3), the data were plotted as bars
(actual ranges and means) on time graphs and the major facts of the
annual cycle of this species were ready for interpretation. Reference
to these graphs ( figs. 1 and 2 ) will be made repeatedly in connection
with the elaboration of the life cycle which follows.

THE LIFE CYCLE

Females. — The maturing females (fig. 1, series A, B, C . . . P)
increased significantly in size from February to early June at which
time they reached a growth plateau that was maintained through July.
Between July and September another significant increase in growth
was evident. Since ovigerous females were first seen in September
this last increase in size is interpreted as representing the "maturity
molt" of the spawning population of females and probably compares
to the male maturity molt {i.e., transformation from second- to first-
form) which must occur prior to copulation. From September to
December the mature females did not show any additional significant
change in size.

The December adult females (group "N") appeared to have in-
creased again in January forming part of group "P", indicating that
at least one molt had taken place. That females molt after the young
leave them was substantiated by laboratory observations. In terms of
the life history this molt may be interpreted as marking the end of
the active reproductive season. Tack (1941) with Orconectes im-
munis and Penn (1943) with Procambarus clarkii both found that
females molted after their young had left, although not immediately.

Group I (fig. 1, series I-IV) is assumed to have been a continuation
of a 1949 group comparable to group "f" of January 1951, and was
probably about a year old. Although not represented by large num-
bers, this series was definitely represented in the samples through
June, but by September seemed to have disappeared, reappearing
presumably in December and January to form the upper ranges of
groups "N" and "P".

In view of the facts that representatives of Series I-IV were not in
the samples from July to December and that several large females
were collected in September showing signs of having already com-
pleted spawning {i.e., with remains of hatchlings' exuviae on their
pleopods) it is thought that other individuals also had been ovigerous
before September and had been overlooked because they were already
in burrows.

Individuals of the series "Q" through "W" were not evident in the
samples from April on, and it is assumed that they had died by this
date. It is assumed that group "P", representing the terminus of two
combined series (A-P and I-IV) which were one and two years old

No. 7

Smith: Life History of Orconectes clypeatus

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88 Tulane Studies in Zoology Vol. 1

respectively, followed the same trends as groups "Q" thru "W" and
died in the spring of 1951.

The presence of various groups of juveniles (fig. 1, series a, b, c . . .
f) from October through January indicates that egg maturation and
embryonic development of the entire population was spread over a
period of several months. The growth trends of each of these groups
cannot be clearly understood from the analyses. The latest juveniles
added to the free-swimming population (group "p") probably repre-
sented the offspring of the few late-spawning females found in Decem-
ber with young still attached to their pleopods. It is assumed that the
January juvenile groups eventually gave rise to groups comparable to
groups "a" and "A" of February 1950.

Males. — As might be expected the growth of the male population
as a whole paralleled that of the females in most respects, but the
interpretation is complicated by the shifts in groupings accompanying
the maturation molt from second- to first-form. The maturing second-
form males (fig. 2, series A, B, C . . . P) increased significantly from
February to May; from May through July they endured a growth
plateau as did the maturing females of the same time period. Some-
time between May and July some of the larger second-form males
apparently matured and molted to first-form, and there is evidence
that the greater percentage actually transformed by May. The period
from February to May was characterized by significant growth in-
creases, and, if the male population exactly paralleled the female a
maturity molt should not have occurred during this time. However,
with the advent of the summer growth plateau and because, of the
overlap of size groups of second- and first-form males in May and
June (fig. 2, groups "C" with "3", and "D" with "4") it is inferred that
some of the larger second-form males had molted into first-form by
May (fig. 2, group 3), and that others molted and became parts of
the groups "4", "5" and "6", and most of the groups "7" and "9" when
the climax of breeding was reached in September.

The majority of group "9" is thought to have continued as first-
form males to December. Groups "H" through "M" represent either
second-form males that had already molted back to first-form (i.e.,
from group "3" through "7") and/or those which would molt to
first-form later in January. From the statistics comparing these groups
the latter assumption seems most justified.

Groups "11", "13" and "14" represent, at least in part, a continu-
ation of group "9". By December and January, individuals of these
groups molted back to the second-form and constituted the upper
parts of groups "N" and "P". Those individuals of groups "N" and
"P" which had not yet molted to first-form apparently did so in
January and constituted the upper part of group "16".

The fact that first-form males molt back to second-form was estab-
lished by laboratory observations of first-form males of the December
sample, most of which had molted back to second-form by January 27th.

Group "e" (fig. 2, series a, b, c . . . e) appears to have developed
by December into the lower part of group "N". By January the
lower part of group "N" is thought to have molted into first-form and

No. 7

Smith: Life History of Orconectes clypeatits

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Tulane Studies tn Zoology

Vol. 1

9 I.I 1.3 1.5 1.7 1.9 2.1

J .3 .9 .7 M 1.1 1.3 13 17

.4 6 8 1.0 1.2 1.4 16 IS Z.0 2.2 .2 .4 .6 .8 1.0 1.2 1.4 1.6 1.8 2.0 2.2

DIAMETER OF EGGS IN MILLIMETERS

Figure 3. Maturation of ovarian eggs. Each sample is represented
b> a percentage frequency histogram in which each bar represents the
percentage of the total number of individuals within the histogram.

then constituted at least the lower part of group "16".

Group I of February (fig. 2, series I-VI) apparently was constituted
of contributions from group "R" (its upper range) and from a group
comparable to group "f" of January (its lower range). Representa-
tives of this series were in each sample from February through July
15th. Prior to February some individuals molted to first-form
(group "1") and from February through July the majority also molted
to first-form contributing to the groups "S" and "2" through "7".
From September through December the remaining few molted to
first-form and were represented by groups "10", "12" and "15".

Groups "Q" and "W" and the upper part of "S" apparently died in

No. 7 Smith: Life History of Orconectes clypeatus 91

the spring as indicated by the absence of the proper size in the May
and June samples. The upper range of group "R" was probably com-
posed of males recently molted from first- to second-form. Group
"T" was a continuation of this group which apparently died by May.

From these conclusions the fate of the January adult group may be
stated. The upper ranges of groups "P" and "16" should have died
first and the remainder of these groups probably died before the next
major reproductive period in 1951. Group "f" should have formed a
group comparable to group "I" of February and lived to be about two
years old.

The growth trends of the male juveniles (fig. 2, series n, o, p . . . t)
were assumed to be essentially similar to those of the female juveniles.

Ovarian Egg Development. — The trend of ovarian egg development
for the year 1950 was determined quantitatively from a comparative
study of percentage frequency histograms (fig. 3) made for each
sample, in which the measurements of ovarian eggs were grouped into
0.10 mm size classes.

Prior to July 12th the ovarian eggs seemed to have been static in
size and in this stage did not show any tendency toward maturing at
different rates as might have been expected from the study of the egg-
laying and rates of development of the young.

Except for the samples of February and June 13th the upper ranges
of these did not exceed 0.6 mm (fig. 3). Only two percent of the
February sample fell within the higher range, and since it was con-
cluded previously that most of the females had already undergone a
molt terminating the major reproductive period, it is inferred that
these larger eggs in February were eventually resorbed. Penn (1943)
for Procambarus clarkii gave evidence that a greater number of eggs
may be developed in the ovary than are laid, and Stephens (1952)
showed that ovarian eggs are readily resorbed by female Orconectes
virilis under experimental conditions.

By September the majority of the eggs were in the large size group
with peaks at 1.3, 1.7 and 2.0 mm. The presence of these large eggs
within the ovaries coincided with the presence of eggs on the pleo-
pods. Within the range below 1.0 mm the peak of 0.6 mm occurred.
It is thought that these eggs eventually matured and were laid.

In September another small peak was noted at the 0.10 mm size.
These are believed to represent recently developed ovarian eggs of
females which had already spawned prior to the date of the sample.

In October females carrying eggs were still present in the samples.
Although no ovarian study was made for this month, due to the in-
adequacy of the sample, it is assumed that the results of the October
1949 analysis would roughly apply. Although apparently egg-laying
was still in progress, the majority of the mature female population
had already spawned, as indicated by the higher percentages of small
eggs.

The December samples showed that at least ninety percent of the
population had already laid eggs prior to this date. The large eggs
still present in the ovaries represented either those that were eventually
resorbed, or that were eventually laid at the individual erratic spawn-

92 Tulane Studies in Zoology Vol. 1

ing times. In general composition this sample compared with that of
February 1950.

By January 1951 the ovarian eggs of the females were all immature,
indicating that the reproductive period for the population as a whole
had definitely terminated.

The reproductive potential was determined from a total count of
the ovarian eggs of all mature non-ovigerous females (nineteen) of
the September sample. The average of these counts is 14.2 eggs per
female within the relatively wide extremes of 9 and 21 (Table 4).

Sex Ratio and Burrowing Responses. — Although burrowing during
drought was adequately confirmed by field observation, these activities
could not be observed directly during the wet periods and an analysis
of the sex ratio of these collections was relied on to furnish these
data. It was assumed that if the water samples, which were apparently
representative of the water population, did not contain the expected
sex ratio, then a significant absence or predominance of one or the
other sex for any particular sample would indicate a burrowing
component.

The "theoretical sex ratio" was based on the actual sex ratio of the
younger male and female homogeneous groups of the February 1950
sample (figs. 1 and 2, groups "A"). The sex ratio for this group was
females/males:=333/310 or 1.07. This group was chosen because
the specimens were large enough to sex easily and all of the young
population was assumed to be in the water at this time of the year.
To determine the chance differences and real differences between this
ratio and the sex ratio of any other group or sample, the limits of a
continuum of likely hypothesis (differences due to chance) and the
limits marking off two continua of unlikely hypothesis (real differ-
ences) were established from the percentage of males to females in
the young February groups.

The results of the comparison (Table 5) show that the females
were not significantly different from the theoretical sex ratio during
the months of February and December 1950 and January 1951, and
possibly April 1950. For these months either both sexes were bur-
rowing or both were not and the fact that both sexes were responding
alike obscures the issue. However, since all these samples were large
in numbers and taken from the ditch during decidedly wet periods
(Table 1) it is assumed that most of the population was probably
in the water.

Those collections in which the percentages of females were sig-
nificantly less than the theoretical ratio were the samples of October
1949 and June 13, July 15 and September 1950. Since the females
were not in the water samples it is assumed that they were burrowing.
Since burrowing coincided in September with the presence of large
eggs in the ovaries (fig. 3) or the presence of eggs on the pleopods,
it is concluded that the egg-laying activity was a stimulus to burrow-
ing. This conclusion lends corroboration to the possibility of erratic
or pre-seasonal reproductive activities of the June 13 and July 15
samples as already indicated (fig. 3) from the study of ovarian eggs.

Analyses of the samples taken from burrows indicates that with one

No. 7

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94 Tulane Studies in Zoology Vol. 1

exception (July 12) there was a significant absence of females in
each of the dry samples. Since it has been established that the entire
population burrowed during drought there is no logical reason why
the females should not have been collected except that the technic
of dry sampling was inadequate. The only other conclusion would
be that of a mass death of only the females during drought times
and this is biologically unacceptable.

SUMMARY AND DISCUSSION OF LIFE CYCLE

During the dry periods the crawfishes burrowed along the sides of
and in the bottom of the ditch instead of migrating to the adjacent
slough, a permanent body of water. There was no evidence during
the course of this study to indicate that the crawfish ever migrated
over dry land; however, dispersal was possible and evidences of it
were observed during periods of flooding rains. Ovigerous females
were found in burrows regardless of the conditions in the ditch.
Indeed, the first ovigerous females collected were dug from burrows
alongside the flooded ditch in September. It may be significant that
immediately following this observation a drought period ensued and
the remainder of the population was forced to burrow. The females
carrying the vulnerable eggs were thus, by natural instincts, already
in protected burrows when the drought conditions began. It would
seem that an ovigerous female could not be so agile or successful a
burrower as one unencumbered by eggs. However, it is not known
whether spawning females burrowed before or after laying their eggs.

The eggs of 0. clypeatus are surprisingly large in comparison to
the relatively small size of the females carrying them. This certainly
is correlated with the low average of 14.2 eggs per female, and per-
haps the whole concept is correlated with the burrow habitat of the
ovigerous females. It is interesting, and possibly significant, that a
very low reproductive potential also obtains in Procambarus hageni-
anns (Faxon) in which the females similarly shun free-water when
ovigerous (Lyle, 1938).

Ovigerous females were collected on September 29 and October
15, 1950. No collections at all were made in November, but females
collected on December 1st were carrying young still clinging to their
pleopods. Although the length of time between oviposition and
hatching, and the time that females normally carry their young was
not determined it is assumed that both time periods were of short
duration since free-living young first appeared in the population in
the October 15th sample, about two weeks after the first ovigerous
females were collected.

The juveniles of January 1951 were represented by five overlapping
age groups. Three groups comparable to the three larger groups of
this sample probably participated in the major spawning period
(September-October) of 1950. Groups comparable to the two younger
groups of January 1951 may have given rise to groups that spawned
erratically (?>., post-seasonal: November -December) during 1950.

As a group the maturing females increased significantly in size
from February to early June. From this date until July 15th they

No. 7

Smith: Life History of Orconectes clypeatus

TABLE

Sex Ratios

Date

Range

Total
Crawfish

Males

Females

Percent of
Females

1949:
Oct. 16

All

1950:
Feb. 24

Adults
Juveniles

122
654

55
316

67
338

55
52

Apr. 9

All

453

240

213

May 12

All

192

107

June 2

All

137

June 13

All

340 .

184

156

July 12

All

July 15

All

169

Sep. 29

All

131

Dec. 29

1951:
Jan. 24

Adults
Juveniles

47
300

107

848

24
146

389

154

459

49
51

remained nearly static in size, but between mid-July and September
they again increased significantly in size, presumably undergoing a
maturity molt. Another molting period ensued during December
and January marking the termination of the active reproductive period
for the majority of the mature females. Apparently most of these
females lived about a year and a half, dying before the next major
spawning period. The females that were already a year old in early
1950 probably constituted an erratic pre-seasonal (July- August)
spawning group in 1950 and died in the first months of 1951 at an
age of about two years.

As a group the males were similar to the females in growth patterns.
The maturing second-form males increased significantly in size until
May at which time the growth plateau was reached. During May a
few of this group molted into first-form males. By September the
majority of them molted into first-form except the very youngest.
These latter apparently transformed towards the end of December
and formed an erratic post-seasonal spawning group among the males.
Many of the first-form males molted back to second-form during
December and January. The participants of the major spawning group
lived to be about one year old, while the participants of the post-
seasonal spawning period lived to be about two years old.

96 Tulane Studies in Zoology Vol. 1

ACKNOWLEDGEMENTS
The writer is deeply indebted to Dr. George Henry Penn under
whose direction this study was made. She is particularly grateful for
his help on many of the field trips and especially for the collections of
June and July which he made; for his many helpful suggestions during
the laboratory exploration, and for his guidance and encouragement
in the presentation of this paper. She is also indebted to Prof. Joseph
A. Ewan, of the Department of Botany of Tulane University, for the
identification of the plants in the ditch, and to Messrs. Lothar E.
Hornuff, Jr. and Edward N. Lambremont for their assistance on sev-
eral field trips.

REFERENCES CITED

Bovbjerg, Richard V. 1952. Comparative ecology and physiology of
the crayfish Orconectes propinquus and Cambarus fodiens.
Physiol Zool, 25 (1) : 34-56.

Cazier, Mont A. and Annette L. Brown 1949. Introduction to quan-
titative systematics. Bull. Amer. Mus. Nat. Hist., 93 (5) : 343-388.

Creaser, Edwin P. 1933a. Seasonal changes in the male population
of Faxonius propinquus (Girard). Occ. Pap. Mus. Zool. Univ.
Mich., No. 253: 1-9.

1933b. Descriptions of some new and poorly known

species of North American crayfishes. Occ. Pap. Mus. Zool. Univ.
Mich., No. 275: 1-21.

Creaser, E. P. and A. I. Ortenburger 1933. The decapod crustaceans
of Oklahoma. Publ. Univ. Okla. Biol. Surv., 5 (2) : 14-47.

Faxon, Walter 1914. Notes on the crayfishes in the United States
National' Museum and the Museum of Comparative Zoology with
descriptions of new species to which is appended a catalogue of
the known species and subspecies. Mem. Mus. Comp. Zool., 40
(8) : 347-427.

Hay, W. P. 1899. Description of two new species of crayfish. Proc.
U. S. Nat. Mus., 22: 121-123.

Hobbs, Horton H., Jr. 1942. The crayfishes of Florida. Univ. Fla.
Publ., Biol. Sci. Ser., 3 (2) : 1-179.

Lyle, Clay 1938. The crawfishes of Mississippi, with special refer-
ence to the biology and control of destructive species. (Abstract).
Iowa State Coll. Jour. Set, 13 (1) : 75-77.

Peatman, John G. 1947. Descriptive and Samjrting Statistics. New
York: Harper and Bros., pp. 1-577.

Penn, George Henry 1942. Observations on the biology of the dwarf
crawfish, Cambarellus shufeldtii (Faxon). Amer. Midi. Nat.,
28 (3) : 644-647.

1943. A study of the life history of the Louisiana

red-crawfish, Cambarus clarkii Girard. Ecology, 24 (1) : 1-18.

1952. The genus Orconectes in Louisiana. Amer.

Midi. Nat., 47 (3) : 743-748.

Stephens, Gwen J. 1952. Mechanisms regulating the reproductive
cycle in the crayfish, Cambarus, I. The female cycle. Physiol.
Zool., 25 (1) : 70-84.

Tack, Peter Isaac 1941. The life history and ecology of the crayfish
Cambarus i in munis Hagen. Amer. Midi. Nat., 25 (2) : 420-446.

Van Deventer, William C. 1937. Studies on the biology of the cray-
fish Cambarus propinqmis Girard. Illinois Biol. Monogr., 15 (3) :
1-67.

■ \\\\. c^iearYSj

'jFUiL&sfs gunman

3S9 ^®(DIL(D@^

Volume 1, Number 8

December 21, 1953

OSTRINCOLA GRACILIS C. B. WILSON, A PARASITE OF
MARINE PELECYPODS IN LOUISIANA

(COPEPODA, CYCLOPOIDA)

ARTHUR G. HUMES,

DEPARTMENT OF BIOLOGY, BOSTON UNIVERSITY,
BOSTON, MASSACHUSETTS.

KSUS. CGm». Z08L
LIBRARY

4 1954

.
'JTY

TULANE UNIVERSITY
NEW ORLEANS

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Trevor Kincaid, Emeritus Professor of Zoology, University of
Washington, Seattle, Washington.

Robert W. Pennak, Professor of Zoology, University of Colo-
rado, Boulder, Colorado.

HARRY C. YEATMAN, Assistant Professor of Biology, University
of the South, Sewanee, Tennessee.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
torial matters, and orders for individual numbers to the editor. Re-
mittances should be made payable to Tulane University.

When citing this series authors are requested to use the following
abbreviations: Tulane Stud* Zool.

Price for this number: $0.25.

George Henry Perm, Editor,
c/o Department of Zoology,
Tulane University,
New Orleans, U. S. A.

wis. zm?. Z80L

UBUklf

JAN 4 1954

OSTRINCOLA GRACIUSJZ.J&-$tii£eN, A PARASITE OF
MARINE PELECYPODS IN LOUISIANA

(COPEPODA, CYCLOPOIDA)
ARTHUR G. HUMES,1

Department of Biology, Boston University,
Boston, Massachusetts.

In June, 1947, a parasitic copepod, Ostrincola gracilis C. B. Wilson
(1944), was discovered in the mantle cavity of four species of pelecy-
pods from the Barataria Bay region of Louisiana. The hosts were the
Virginia oyster (Crassostrea virginica Gmelin), the ribbed mussel
{Modiolus demissus granosissimus Sowerby), the recurved mussel
(Mytilus recurvus Rafinesque), and the hard shelled clam or quahaug
{Venus mercenaria mercenaria Linne.).

A study of the copepods thus obtained has revealed certain im-
portant omissions and inaccuracies in the original description. The
holotype and allotype specimens, U. S. National Museum No. 79697,
have not been available for dissection and study, but a male and a
female, U. S. National Museum No. 79698, having the same collection
data as the holotype and allotype and apparently paratypes though
not designated as such by Wilson, have been compared in detail with
the Louisiana forms. The specimens from Louisiana proved to be
conspecific with the actual specimens of 0. gracilis as identified by
Wilson.

For helpful suggestions in connection with this paper I am indebted
to Mrs. Mildred S. Wilson.

Female. — The body ( fig. 1 ) in living specimens is colorless,
except for a dark reddish-black median eye and a brownish intestine.
(In glycerine and polyvinyl alcohol mounts the eye is bluish.) The
total length (from the tip of the head to the posterior end of the
caudal rami), based on an average of five specimens, is 1.083 mm and
the greatest width of the cephalothorax is 0.274 mm. The abdomen is
three-segmented. The ventral surface of the genital segment and
abdomen bears minute spines arranged as in figure 2, but the dorsal
surface lacks spines. The caudal ramus (fig. 3) bears three terminal
setae and three along its outer edge. The terminal setae in many
specimens show brownish accretions around their tips (fig. 4).
The egg sacs (fig. 5) both contain about seven eggs and are
attached dorsolaterally. The first antenna (fig. 6) has seven podo-
meres and bears several aesthetes. The arrangement of the setae
on the distal podomere suggests a subdivision into two podomeres

1 The material upon which this paper is based was collected while
the author was a guest investigator at the Louisiana State University
Marine Laboratory at Grand Isle, Louisiana.

100

Tulane Studies in Zoology

Vol. 1

No. 8 Humes: Parasite of Marine Pelecypods 101

but there is no articulation. The second antenna (fig. 7) has a
notch-like interruption on the inner margin of the claw (fig. 8).

The mouthparts (fig. 9) are much reduced. The labrum is wider
than long and bears on each side a group of small spines. Its posterior
edge seems to be entire and not deeply incised, although this feature
is difficult to ascertain because of the impinging of the labrum and
the mouthparts. The mandible (fig. 10) has a somewhat elongated
base with an apical armature of four elements. The first maxilla (fig.
11) is minute and arises from' a point along the distal half of the
basal part of the mandible. In unmounted specimens the first maxillae
extend nearly vertically from the ventral body surface, but in pressed
whole mounts they appear to be directed more posteriorly. Each
first maxilla consists of a single lobe bearing four setae which are
progressively shorter toward the midline of the body. The second
maxilla (fig. 12) has a much inflated basal part and a relatively
small apex with tripartite armature. The distal portions of both the
mandibles and the second maxillae in unmounted specimens are
directed vertically from the ventral body surface and somewhat an-
teriorly. The mandibles and both pairs of maxillae are minute and
closely appressed, so that it is only with great care that they can be
isolated for study. The labium is small and rather poorly delimited,
with a longitudinal concavity on its posterior surface. The maxilli-
peds are lacking. In the median region of the head posterior to the
labium there is a prominent postoral protuberance, especially con-
spicuous in lateral view. In the midline of the body on the anterior
surface of the ridge between the bases of the first pair of legs there
is a trilobed structure (fig. 13). In pressed specimens this trilobed
area appears to be overlapped by the posterior edge of the postoral
protuberance.

The first, second, third, and fourth swimming legs are shown in
figures 14-17 respectively. The spine and setal formula is as follows:

legl

exp end

leg 2
exp end

leg 3

exp end

leg 4
exp end

first podomere
second podomere
third podomere

1:0 0:1

1:1 0:1

8 6

1:0 0:1

1:1 0:2

9 6

1:0 0:1

1:1 0:2

8 6

1:0 0:1

1:1 0:2

8 5

The fifth leg (fig. 18) has two podomeres, the proximal
with a single outer seta, the distal one expanded into a

one small
broad, in-

Figures 1-9. Ostrincola gracilis, female: 1, dorsal view; 2, genital
segment and abdomen, ventral view; 3, caudal ramus; 4, terminal
setae of caudal ramus with accretions; 5, egg sac; 6, first antenna;
7, second antenna; 8, detail of claw of second antenna; 9, area of
mouthparts.

102

Tulane Studies in Zoology

Vol. 1

No. 8 Humes: Parasite of Marine Pelecypods 103

wardly concave disk with four marginal setae.

Male. — The color in life resembles that of the female. The total
length, based on an average of five specimens, is 0.760 mm and the
greatest width of the cephalothorax is 0.200 mm. The abdomen is
four-segmented (fig. 19). The ventral surface of the genital seg-
ment and abdomen bears minute spines as in figure 20. The caudal
rami are like those of the female. The first antenna (fig. 21) has
seven podomeres, with all but the proximal one having an aesthete.

The second antennae, mandibles, first maxillae, and second maxillae
are like those of the female. The maxilliped (figs. 22, 23) has
five podomeres, the proximal one short, the second large and spinose
on its outer and inner margins, the third short, the fourth short also
with an inner seta, and the last elongated, arcuate, and slender. The
last podomere is as long as or slightly longer than the rest of the
podomeres together.

The four pairs of swimming legs are identical in structure with
those of the female, having the same spine and setal arrangement.
The fifth leg (fig. 24) has two podomeres, the first with a single
outer seta, the second longer, not expanded, but with more or less
parallel margins, and with four marginal setae. The sixth leg (figs.
20, 25) consists of two setae at the posterior corner of the genital
segment. In a few males some of the spines near the . sixth legs
show brownish caps covering their tips (fig. 26).

Remarks. — Ostrincola seems to stand between the ergasilid genus
Myicola Wright (1885) and the Lichomolgidae as represented by
the genus Pseudomyicola Yamaguti (1936). The segmentation and
armature of the first antenna, the mandible, the first and second
maxilla, the male maxilliped, and the segmentation and armature of
legs 1-4 resemble Myicola metisiensis Wright. In two respects at
least Ostrincola differs from Myicola. The second antenna is rather
elongated and slender, while in Myicola it is shortened and robust.
The distal podomere of the fifth leg in the female is much expanded,
while in Myicola it has nearly subparallel margins. In both these
characters Ostrincola resembles the condition in Pseudomyicola, differ-
ing from that genus, however, in the first antenna, the mandible, and
other features.

The generic validity of Ostrincola seems to rest upon the future
discovery of other related species. In the meantime, however, the
original generic description may be emended as follows:

Figures 10-16, 18. Ostrincola gracilis, female: 10, mandible; 11,
first maxilla; 12, apex of second maxilla; 13, trilobed structure be-
tween bases of first legs; 14, first leg; 15, second leg; 16, third leg;
18, fifth leg.

104

Tulane Studies in Zoology

Vol. 1

No. 8 Humes: Parasite of Marine Pelecypods 105

OSTRINCOLA Wilson 1944, emended diagnosis

Body cyclopoid with head and first thoracic segments fused. Tho-
racic segments 2-5 diminishing regularly in width. Genital segment
longer than wide. Abdomen three-segmented in the female, four-
segmented in the male. Caudal rami much elongated and sub-
cylindrical. Sexual dimorphism not pronounced. Attachments of
egg sacs dorsolateral; eggs relatively few in number.

First antenna in both sexes with seven podomeres. Second antenna
uniramous, with four podomeres, the penultimate relatively long and
slender and the last a curved claw. Mandible with a moderately long
basal portion bearing an apical armature of four parts, directed an-
teriorly. First maxilla a minute somewhat posteriorly directed lobe
arising from the area immediately behind the distal half of the base
of the mandible and bearing four unequal setae. Second maxilla with
a much inflated basal part and a small apical portion with tripartite
armature, directed anteriorly. Anterior to the mouthparts a swollen
labrum without a deeply incised posterior edge, and posterior to them
a rather weakly developed labium. Behind the labium a swollen
postoral protuberance. Maxillipeds lacking in the female. The male
maxilliped large, with five podomeres, the second podomere elongated
and relatively stout, the last podomere slender, arcuate, and about as
long as the other four podomeres together.

Legs 1-4 biramous, the rami with three podomeres, and the spine
and setal formula as in Myicola. Fifth leg uniramous in both sexes,
with two podomeres; the distal podomere in the female expanded into
a broad, inwardly concave disk armed along the margin with four
setae, in the male elongated with subparallel margins, similarly armed.

Genotype. — Ostrincola gracilis Wilson 1944.

The major points in the emendation concern the number of ab-
dominal somites, the number of first antennal podomeres, the nature
of the mouthparts, and the number of podomeres in the fifth legs.

Occurrence. — In Barataria Bay 0. gracilis was found near Chene
Fleurie in Mytilus and Crassostrea, in Sugarhouse Bend Bayou at
Grand Terre in Mytilus and Crassostrea, and in the tidal marsh back
of Grand Isle in Venus, Crassostrea, and Modiolus. The collection
data are summarized in Table 1.

Figures 17, 19-26. Ostrincola gracilis, female: 17, fourth leg. Male:
19, dorsal view; 20, genital segment and abdomen, ventral view; 21,
first antenna; 22, area of mouthparts; 23, maxilliped; 24, fifth leg;
25, sixth leg; 26, setae of sixth leg with cap-like coverings.

106 Tulane Studies in Zoology Vol. 1

TABLE 1

Incidence of Ostrincola gracilis in pelecypods of the Barataria

Bay region of Louisiana.

Clam Host

Number
Examined

Number

with

0. gracilis

Percent
Parasitized

Average
Number
Per Host

Crassostrea virginica

399

157

39.3

1.88 (1-11)

Modiolus demissus
granosissimus

241

36.9

1.73 (1-7)

Mytilus recurvus

15.5

1.14 (1-2)

Venus mercenaria
mercenaria

100

686

254

37.0

1.81 (1-11)

Ecological notes concerning the four species parasitized are to be
found in the publication of Harry (1942). According to observa-
tions previously made (Humes, 1942) the water in Barataria Bay
seldom exceeds ten feet in depth, except in the passes leading to the
open Gulf. The bottom is largely composed of sand and silt with
extensive areas devoted to oyster beds. The summer water temperature
ranges from 25.8 to 29.8° C. The salinity at Chene Fleurie is 9-18
grams per liter, while at the eastern end of Grand Terre near Sugar-
house Bend Bayou it is 25.30. Although no ecological information was
given by Wilson (1944) regarding the type specimens at Beaufort,
North Carolina, it seems probable on the basis of the Louisiana
material that this copepod will be found as a parasite in pelecypods
living in shallow-water, estuarine regions which are protected from
the open sea.

Explanation of Figures

All figures were drawn with the aid of a camera lucida. Scale A
applies to figures 1 and 19; scale B to figures 2, 5, and 20; scale C to
figures 3, 4, 13, 23, 25, and 26; scale D to figures 6, 7, 14-18, 21, and
24; scale E to figures 8, and 10-12; and, scale F to figures 9 and 22.

REFERENCES CITED

Harry, Harold W. 1942. List of Mollusca of Grand Isle, Louisiana,
recorded from the Louisiana State University Marine Laboratory,
1929-1941. Occ. Pap. Marine Lab., La. State Univ., No. 1: 1-13.

Humes, Arthur G. 1942. The morphology, taxonomy, and bionomics
of the nemertean genus Carcinonemertes. Illinois Biol Monogr.,
18 (4) : 1-105.

Wilson, C. B'. 1944. Parasitic copepods in the United States Na-
tional Museum. Proc. U. S. Nat. Mus., 94: 529-582.

No. 8 Humes: Parasite of Marine Pelecypods 107

Wright, R. R. 1885. On a parasitic copepod of the clam. Amer. Nat.,
19 (2) : 118-124.

Yamaguti, S. 1936. Parasitic copepods from mollusks of Japan, I.
Jap. Jour. Zool, 7 (1) : 113-127.

\l » ' ^

■w • **-..>' _

Volume 1, Number 9

February 8, 1954

HYBRID INVIABILITY BETWEEN RANA PIPIENS FROM
WISCONSIN AND MEXICO

E. PETER VOLPE,

DEPARTMENT OF BI0L00Y3 NEWOOMB COLLEGE, TULANB
UNIVERSITY, NEW ORLEANS, LOUISIANA

FEB

1 8 1954

TULANE UNIVERSITY
NEW ORLEANS

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Victor C. Twitty, Professor of Zoology, Stanford University,

Stanford University, California.
John A. Moore, Professor of Zoology, Columbia University,

New York, New York.
Albert P. Blair, Associate Professor of Zoology, University of

Tulsa, Tulsa, Oklahoma.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
torial matters, and orders for individual numbers to the editor. Re-
mittances should be made payable to Tulane University.

When citing this series authors are requested to use the following
abbreviations: Tulane Stud. Zool.

Price for this number: $0.35.

George Henry Penn, Editor,
c/o Department of Zoology,
Tulane University,
New Orleans, U. S. A.

FEB 1 8 1954

HYBRID INVIABILITY BETWEEN RANA PIP1ENS FROM
WISCONSIN AND MEXICO1

E. PETER VOLPE,

Department of Biology, Neivcomb College, Tulane
University, New Orleans, Louisiana

Laboratory crosses among individuals of various populations of the
widely-distributed meadow frog, Rana pipiens, have revealed the ex-
tent to which intraspecific gene exchange is possible. The occurrence
of abnormalities in hybrids between certain geographical populations
of Rs pipiens has been demonstrated by Moore (1949a). In general,
the intensity of hybrid defects is inversely proportional to the north-
south distance between the localities of the parents employed in the
laboratory hybridizations. It has also been shown (Moore, 1949b)
that the geographically extreme northern and southern populations
of this species differ in embryonic rate of development and embryonic
temperature tolerance. Moore has inferred that these embryological
differences represent differential adaptations to environmental tem-
peratures, and he further surmises that the embryonic defects in
hybrids between northern and southern individuals may be associated
with the different temperature adaptations of the parental eggs.

The most severe abnormalities have been observed in hybrids be-
tween R. pipiens from Alburg, Vermont and two localities in eastern
Mexico, Monterrey in Neuvo Leon and Axtla in San Luis Potosi
(Moore, 1947).

In the present investigation, meadow frogs obtained simultaneously
from a different northern locality, Oshkosh, Wisconsin and several
localities in the State of Tamaulipas, Mexico provided the opportunity
to hybridize the respective groups, and thus contribute additional
information on the problem of geographical differentiation and the
development of isolating mechanisms in populations of R. pipiens*

Experimental
A comparison was made of the rate of embryonic development
and embryonic temperature tolerance of R. pipiens from Mexico and
Wisconsin to determine if differences existed, and reciprocal hybrid
crosses were conducted to ascertain the extent of developmental com-
patibility. The methods of experimentation were essentially similar
to those employed in other work of this type (Moore, 1946, 1949b;
Volpe, 1952, 1953). The technique, and the modifications introduced,
will be brought out during the discussion of the experiments.

Embryonic temperature adaptations. — Ovulation of Mexican and
Wisconsin meadow frogs was induced by pituitary injection. The

1 Aided by a grant from the Tulane University Council on Re-
search. I am indebted to Dr. R. M. Darnell of Tulane University for
collecting the meadow frogs from Mexico, and to Miss Vera Morel for
the drawings used in illustrating this paper.

112 Tulane Studies in Zoology Vol. 1

eggs were stripped from the females into sperm suspensions prepared
by macerating the testes of a male in 10-20 cc of 0.1% Ringer's
solution. Eight artificial fertilizations, in groups of two, were con-
ducted. Each fertilization of a Mexican 9 X Mexican $ was carried
out simultaneously with a cross Wisconsin 2 X Wisconsin S. By
conducting parallel fertilizations, differences in rate of development
of Mexican and Wisconsin embryos were readily apparent. The four
ovulated Mexican females were derived from four different localities,
as follows: (1) Storm's Ranch, 7 km. northeast of Gomez Farias,
Tamaulipas; (2) La Union, 9V^> km. north of Gomez Farias, Tam-
aulipas; (3) El-Mante-Tampico Highway, 29 km. east of El Mante,
Tamaulipas; (4) El Mante-Tampico Highway, 72 km. east of El
Mante, Tamaulipas. All four Wisconsin mating pairs employed came
from Oshkosh, Wisconsin.

The egg masses from any single fertilization were cut into small
clusters of 9 to 13 eggs to permit more surface for respiration. The
eggs clusters were placed in finger bowls, each finger bowl contain-
ing ten clusters (referred to as a "group") in 200 cc of 0.1%
Ringer's solution. The finger bowls were than distributed to con-
stant temperature units, which maintained temperatures accurate to
±0.2 °C. Development of four groups of Wisconsin eggs and four
groups of Mexican eggs was observed at 18.9°C, 21.1 °C, 27.5 °C,
29.5 °C, and 33 °C.

The eggs derived from the Mexican females were larger than
those obtained from the Wisconsin females. Measurements of the
eggs (vitellus diameter) prior to first cleavage were made by means
of an ocular micrometer calibrated from a 1/100 millimeter-ruled
stage micrometer. The diameters of 86 Wisconsin eggs ranged from
1.60 mm. to 1.95 mm., the mean and standard deviation being
1.73 ±0.08. Similar measurements of 101 Mexican eggs indicated
a range of 1.95 mm. to 2.20 mm. (2.10±0.06). The difference
is significant (the difference of the means is three times the stand-
ard error of the difference). Although a few of the Wisconsin eggs
approached the size of the Mexican eggs, a cluster of the latter could
be readily distinguished with the unaided eye from a cluster of the
former. This detectable difference in egg size (recall the relation
of the volume of a sphere to its radius) made it possible to keep
both the Mexican and Wisconsin clusters of eggs in the same finger
bowl containing Ringer's solution at a desired temperature. This
procedure reduced the error of developmental deviations due to any
temperature variations which may have been encountered if each
group of eggs had been kept in separate finger bowls. No difficulty
was experienced in identifying the two groups of eggs while they
developed within the jelly membranes. To obviate any difficulty
in later development, the embryos of each group just prior to hatch-
ing were separated into individual finger bowls containing fresh
0.1% Ringer's solution which previously had been brought to the
desired temperature.

No. 9

Volpe: Hybrid lnviability in Rana pipiens

113

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114 Tulane Studies in Zoology Vol. 1

The rate of development was determined by examining the groups
of embryos at each of the constant temperatures at frequent intervals
from first cleavage to gill circulation. Certain problems arise in
obtaining an adequate measure of the time required by a group of
embryos to reach a given stage. Even when special precautions are
taken to provide as uniform an environment as possible, eggs within
a group exhibit variation in the speed with which they reach a given
point in development. Thus, to permit statistical comparison between
two groups of developing eggs, it would be desirable to make re-
peated observations to determine when 100 percent of the eggs had
reached a particular morphological stage. This is laborious and
tedious and even if this procedure were practical, it would be neces-
sary to break the continuous embryonic development into a series
of extremely minute discontinuous steps to permit an accurate de-
termination of the extent of variation. The problem was attacked
by selecting morphological stages at which recognizable specific proc-
esses begin, and the onset of a given stage was taken as the time
at which 50 percent or more of the eggs within a group entered
that stage. The stages drawn up by Pollister and Moore (1937)
were employed, with the further refinement of designating the
stages as early (E), middle (M), and late (L). In comparing two
groups of eggs which were developing at different speeds, an esti-
mate of the variability was obtained by noting the percent of
embryos in the more slowly developing group which were as ad-
vanced as the majority of embryos in the second group which exhibited
the faster rate of development. In all experiments, it was found that
only five percent or less of the more slowly developing group of
embryos were as advanced as the majority (50 percent or more) of
the faster developing group of embryos; nor were there more than
five percent of the faster developing group of embryos in as early
a stage as the majority (50 percent or more) of the more slowly
developing group of embryos. Moreover, the arbitrary criterion of
50 percent was found to be a conservative figure. From fertilization
to stage 17 (tail bud stage) 80 percent or more of the embryos
reached a given stage simultaneously; from stage 18 (muscular con-
traction) onward, 70 percent or more of the embryos entered a
particular stage at the same time. Thus, although it is difficult to
analyse the spectrum of variation, the ascertainment at any one read-
ing of the norm of a group and the extreme variants of that group
(in relation to the other group being compared) permits an accurate
determination of comparative rates of development.

No differences could be detected in the rate of development of the
four groups of Mexican embryos at each of the experimental tem-
peratures. However, the groups of Mexican embryos differed in
rate of development from those derived from Wisconsin. In Table
1 a comparison is made of the time in hours required by 50 percent
or more Mexican and Wisconsin embryos within a group to reach
the experimental end-point, stage 20, at two temperatures, 18.9°C

No. 9 Volpe: Hybrid Inviability in Rana pipiens 115

110-

90-

§70-

o o

— i i • i |

• MEXICO
o WISCONSIN

t — i — i — i — | — i — I — I — l — I — i — i — i — i 1

15 20 25 30 35

TEMPERATURE

Figure 1. A comparison of the mean time, in hours, between stages
3 and 20 at different temperatures, in degrees Centigrade, of embryos
of Rana pipiens from Mexico and Wisconsin.

and 21.1 °C. At 18.9 °C, the embryos from each of the different
localities in Mexico developed more slowly than the Wisconsin em-
bryos. At 21.1 °C (and higher temperatures), the Mexican embryos
developed more rapidly. Figure 1 shows a plot of the mean time
in hours between stages 3 and 20 required by four groups of Mexican
embryos and four groups of Wisconsin embryos at the different
temperatures employed. The faster rate of development of Mexican
embryos at higher temperatures may be associated with the greater
tolerance of these embryos to high temperatures.

To estimate temperature tolerance, records were made of the types
and proportions of embryonic abnormalities at each temperature.
The Mexican embryos are decidedly better adapted to the higher
temperatures. At 33.0°C, early death of all of the Wisconsin em-
bryos was evidenced by irregular cleavage furrows. At this same
temperature, the Mexican eggs cleaved normally, but the majority of
the eggs later exhibited defects in gastrulation. However, an average
of 13 percent for the four groups of embryos (57 of a total of 437
eggs) developed to the gill circulation stage (stage 20). At 29.5 °C,

116

Tulane Studies in Zoology

Vol. 1

.Worma.!

Abnormal

tsiormail

Normal

Uorrrual

Abnormal

jbJormaJ

Abnormal

Figures 2-8. Development of hybrid embryos from the cross La
Union, Mexico 5 X Oshkosh, Wisconsin $. Control embryos (La
Union, Mexico $ X La Union, Mexico $ ) are shown in some figures
for comparison. For full explanations, see text. 2. The pitted ap-
pearance of the ectodermal surface of an early hybrid gastrula;
3. The furrow encircling the hybrid egg and the narrow depression
on the dorsal side representing the neural groove; 4. Top: normal
appearance of the tail-bud stage (stage 17) in a control embryo;
Bottom: hybrid embryo with extreme reduction of head, fused oral
suckers projecting from head, and the ventral abdominal bulge;
5. Top: heart beat stage (stage 19) in a control embryo; Bottom:
shrivelled appearance of hybrid embryo showing abbreviated head,

No. 9 Volpe: Hybrid lnviability in Rang pipiens 117

only a few developmental irregularities (total average of nine percent
for four groups of embryos) were noted in the Mexican embryos.
In contrast, the Wisconsin embryos developed only as far as the
neurula stage. Moreover, development was normal in the Mexican
embryos at 27.5 °C, whereas 85 percent (390 of a total of 460 eggs)
of the Wisconsin tadpoles were distorted.

At the low temperature extreme, adequate temperature control was
not available. However, some indication of the differential effects
of low temperature on the embryos was obtained by placing the
developing eggs in a refrigerator which fluctuated from 6°C to 9°C.
Four groups of Mexican eggs (total of 392 eggs) failed to cleave and
436 Wisconsin eggs developed normally up to stage 17 (300 hours),
at which time the experiment was terminated. It appears that the
embryos of R. pipiens from Wisconsin behave like those of other
northerly distributed amphibians in being able to tolerate low tem-
peratures.

In all probability, the differences in rate of development and
temperature tolerance of Mexican and Wisconsin eggs represent
adjustments that adapt the eggs to their respective environments.
However, data on prevailing environmental conditions, particularly
temperatures of their breeding waters, are lacking. Also, it seems
reasonable to suspect that the improper interadjustments of morpho-
genetic movements observed in hybrid development (discussed below)
are attributable, at least in part, to conflicting modes of action of
temperature-related processes.

Reciprocal artificial hybridizations. — The procedure in artificial
hybridization consisted of fertilizing the eggs of one female from a
particular locality in two batches: the first with the sperm of a male
derived from the same locality ("control"), and the second with the
sperm of a male from a different locality ("hybrid"). Eight hybrid
crosses were conducted, four of which involved a Mexican pipiens
egg and a Wisconsin pipiens sperm each with controls ("Type A"
cross); the others were reciprocal crosses each with controls ("Type
B" cross). The hybrid embryos from the two types of crosses were

ventral abdominal bulge, and the curved tail (the confining inner
jelly membrane is not shown); 6. Top: gill circulation stage (stage
20) in a control embryo which has hatched from the jelly mem-
branes; Bottom: hybrid embryo showing little progress made over
previous stage (fig. 5) ; the hybrid embryo is still trapped within
the inner jelly membrane, but this is not shown in the illustration;
7. Top: gill circulation stage (stage 20) in a control embryo; Bot-
tom: an exceptional hybrid embryo which has hatched, showing the
wrinkled blastocoele roof in the ventral abdominal region and the
microcephalic condition with the downward projecting fused oral
suckers; 8. Top: tail fin circulation stage (stage 22) in a control
embryo; Bottom.: a surviving hybrid embryo in which considerable
reorganization has taken place: the ventral abdominal bulge has
disappeared and the head tends toward a normal condition ; however,
mouth and eye formations are atypical.

118 Tulane Studies tn Zoology VoL 1

characterized by retarded developmental rates and extensive structural
abnormalities. Within each type of cross the results were essentially
similar; thus only a single cross of each type will be described in detail.

Type A, Cross 1:

La Union, Mexico 2 X Oshkosh, Wisconsin $
La Union, Mexico 2 X La Union, Mexico $ (Control)
Conducted at 18.9°C
The rate of development in the early cleavage stages was maternal
and the cleavage furrows were normal. The first indication of hybrid
abnormality appeared during gastrulation. The surface of the pre-
sumptive ectoderm area overlying the blastocoele became wrinkled
and presented a pitted appearance (fig. 2). During the closure of
the blastopore and the subsequent formation of the neural folds, the
series of pits arranged themselves in the form of a groove or furrow
encircling the ventral portion of the egg (fig. 3). The groove
marked externally the floor of the blastocoele. This was determined
by puncturing a few eggs with a needle slightly below the groove.
Blastocoele fluid escaped and yolk cells occupied the internal area
at the level delimited externally by the groove. Thus the blastocoele
failed to become completely obliterated during the gastrular move-
ments. The persistent blastocoele later assumed the shape of a bulge
in the ventral abdominal region of the developing hybrid embryos.
The neural folds that formed in the hybrid embryos were narrower
and closer together than in the control embryos. This condition
foreshadowed the future reduction of the head. Figure 4 shows the
extreme reduction and abnormalities of the anterior region of hybrid
embryos at the tail bud stage. The oral suckers were greatly reduced
in size and fused; the olfactory pits were absent or, if present, ap-
proached one another toward the mid-line; the stomadeal groove
was absent; the gill plate showed no visible differentiation into
arches; and the ventral abdominal bulge filled with blastocoele fluid
was prominent. By late stage 19, the control embryos had already
hatched (fig. 5). The majority of the hybrid embryos remained
curled within the inner jelly membrane and failed to hatch. The
structural abnormalities in the hybrid embryos were pronounced. The
ventral abdominal bulge remained relatively large; the fused oral
suckers projected downward; no stomadeal pit was present; and the
gill plates remained undifferentiated and drawn close to the head.
The hybrid embryos appeared arrested at this time. When the con-
trols reached stage 20 (gill circulation), no advances were made by
the hybrid embryos over the prior stage (fig. 6). Most of them
cytolyzed within the inner jelly mass. However, some of the hybrids
hatched. In those hatching, either the bulge in the ventral ectoderm
became wrinkled and cytolysis ensued, or, in a very small percent, the
bulge became reduced in size (fig. 7). Of the latter a few continued
development into stage 22 (fig. 8). In these surviving hybrid em-
bryos, the small fused suckers began to disappear, but the development
of the mouth structures and eyes were much delayed and small.

No. 9 Volpe: Hybrid Inviability in Rana pipiens H9

These hybrids were transferred to a balanced aquarium, but all failed
to live. Death was associated with reduced heads, abnormal eyes
and atypical mouthparts, as control embryos placed in the same tank
at the same time continued developing.

The hybrid abnormalities described above were also noticed in the
other crosses of Mexican 9 X Wisconsin $ ("Type A"). Hybrid
embryos of this type of cross characteristically exhibited reduced heads
with fused suckers and fused olfactory pits. In each of the four
"Type A" crosses conducted, all hybrid embryos died.

Type B, Cross 1:

Oshkosh, Wisconsin 9 X 29 km. e. El Mante, Mexico $
Oshkosh, Wisconsin 9 X Oshkosh, Wisconsin $ (Control)
Conducted at 18.9 °C

The defects observed in these hybrid embryos were different, in
fact almost opposite to those described in the "Type A" cross. Many
embryos failed to recover from the effects of abnormal gastrulation.
The disintegrating gastrulae exhibited open blastopores with ex-
truding yolk plugs. However, a fair percent formed neural plates
though the blastopore remained open (fig. 9). The neural folds
that developed in the anterior end of the hybrid embryos were more
divergent than encountered in the controls. This foreshadowed the
enlarged head that appeared in later development. As shown in
figure 9, the hybrid embryos were markedly retarded in development
(the controls were in late stage 16). During later development
(fig. 10), the hybrid embryos showed open blastopores of varied
sizes. An atypical tail bud developed from the median dorsal lip
of the open blastopore. Characteristic defects, other than the per-
sistent yolk plug and shortened tail, were the enormous head, the
abbreviated body form, and the unusually immense, widely-separated
oral suckers. When the controls reached stage 19 (fig. 11), the
surviving hybrid embryos were considerably retarded. An unnatural
mucus mass filled the space between the huge oral suckers. As the
controls progressed to stage 20 (fig. 12) the hybrids showed little
developmental advance. The oral suckers remained large and diver-
gent, and the blastopore beneath a shortened tail remained open in
most embryos. By the time the controls were in stage 22 (fig. 13)
all the hybrid embryos died. Cytolysis appeared to begin first in the
yolk cells of the exposed yolk plug. The cytolyzing embryos showed
enlarged heads, immense oral suckers, curved tails, and poorly dif-
ferentiated gill plates. A few, prior to death, showed the earliest
beginnings of external gills, but corpuscles were never observed
coursing through them.

The hybrid embryos in the three other "Type B" crosses were char-
acterized by the same kind and intensity of abnormalities. Not a
single functional tadpole was obtained in all the crosses involving
Wisconsin eggs and Mexican sperm.

120

Tulane Studies in Zoology

VoL 1

UormaJ

Uormal

Jsiorma.1

>Jorm<a.1

Figures 9-13. Development of hybrid embryos from the cross Osh-
kosh, Wisconsin 2 X Tampico, Mexico $. Control embryos (Osh-
kosh, Wisconsin £ X Oshkosh, Wisconsin $ ) are shown in all figures
for comparison. For full explanation, see text. 9. Top: a control
embryo approaching tail bud stage (stage 17) ; Bottom: retarded
hybrid embryo in medullary plate stage (stage 13), showing the
open blastopore with the large, extruding yolk plug; 10. Top: tail
bud stage (stage 17) in a control embryo; Bottom: hybrid embryo
showing the exposed yolk plug and the large, divergent oral suckers;
11. Top: heart beat stage (stage 19) in a control embryo; Bottom:
hybrid embryo with an enlarged head, stubby tail, and a thick mucus
mass projecting from the overactive oral suckers; 12. Top: gill cir-
culation stage (stage 20) in a control embryo; Bottom: a hybrid
embryo arrested in development; abnormal in those respects previ-
ously noted; 13. Top: tail fin circulation stage (stage 22) in a con-
trol embryo; Bottom: appearance of a hybrid embryo just prior to
cytolysis; note the enlarged head, shortened body form, and poorly
differentiated tail.

No. 9 Volpe: Hybrid Inviability in Rana pipiens 121

Discussion

It is clear that isolation through hybrid inviability has reached a
high intensity between Oshkosh, Wisconsin Rana pipiens and those
from the area of Tamaulipas, Mexico. Moore (1947) demonstrated
a similar high degree of incompatibility between Alburg, Vermont
and eastern Mexico meadow frogs. It may have been argued that
the observed hybrid abnormalities were due to certain characteristics
unique only to Vermont meadow frogs among northern pipiens or
to an equally peculiar property of those meadow frogs from the par-
ticular Mexican localities studied by Moore. Similar results in this
investigation utilizing pipiens from another northern locality and
different Mexican localities suggest the likelihood that all northern
meadow frogs could not cross with their southerly-distributed low-
land relatives in Mexico. The geographically extreme members of
the species range have built up different adaptive gene complexes
to the extent that they are incapable of producing viable hybrids in
the laboratory.

It must be admitted that the probability of a Wisconsin meadow
frog crossing with a Mexican frog in nature is extremely remote.
Yet it is just this point which emphasizes the fact that incipient
isolating mechanisms do not develop initially for the effect itself,
but are simply the inevitable consequence of two populations ac-
cumulating sufficient adaptive genetic differences during a period
of geographical separation. Muller (1942) was among the first to
favor this concept that isolating mechanisms arise as a by-product
of genetic divergence of allopatric populations. The genetic changes
which arise to better adapt one population to particular environ-
mental factors may also be instrumental in isolating to varied degrees
that population from other populations which may possess different
adaptive, incidentally isolating, pleiotropic alleles. In our case, genetic
differentiation of the allopatric pipiens populations has proceeded
to such a marked degree that isolation is almost complete. Pre-
sumably the genes or gene complex governing different temperature
requirements of the two types of eggs also have very strong isolating
effects. If the Wisconsin and Mexican pipiens were ever to meet
in nature, natural selection would probably reinforce the isolation
thru hybrid inviability by additional mechanisms which would
guard against the production of hybrids and thus prevent the wastage
of reproductive energy. Recently, Koopman (1950) has demon-
strated experimentally an increase in the amount of reproductive
isolation between two species of Drosophila as a result of continual
artificial selection.

Of embryological interest is the demonstration that the results of
hybridization between the two geographically extreme groups are
very dissimilar in the reciprocal crosses. The gastrulation process
is differentially affected in each type of hybrid development. In

122 Tulane Studies in Zoology Vol. 1

the cross Mexican pipiens 2 X Wisconsin pipiens S, the imagi-
native phase (internal rearrangement of groups of cells) of gastru-
lation is most seriously hampered, whereas in the reciprocal cross
the progress of the blastopore lips over the yolk (epiboly of gastru-
lation) appears to be most disturbed. These early defects in the
gastrular movements are responsible for the major subsequent ab-
normal form changes in each type of cross. In the cross Mexican
pipiens 2 X Wisconsin pipiens $ , restriction of internal yolk move-
ment results in the failure of the blastocoele to be completely oblit-
erated. As the persistent blastocoele offers resistance to the in-
voluting chorda-mesoderm, the latter is abnormally disposed in the
anterior head region and this probably accounts for the subsequent
microcephaly. In the reciprocal cross (Wisconsin pipiens 2 X
Mexican pipiens 8 ) , restriction of the epibolic spreading of the
animal hemisphere results in a permanently open . blastopore. The
exposed yolk-plug later cytolyzes under the irregularly curved, short-
ened tail. Although the abnormalities in the posterior region of
these hybrid embryos are traceable back to an open blastopore, the
origin of macrocephaly in the anterior region is not apparent from
the observations made.

As has been shown, normally fertilized eggs of Wisconsin and
Mexican pipiens differ considerably in their rate of development.
This is the most obvious physiological difference between the two
eggs, and one may be tempted to propose that an asynchronous de-
velopmental rate in the hybrid egg is the factor initially responsible
for the observed abnormalities. It is hazardous, however, to suggest
that this may be the actual causative mechanism. The hybridization
experiments were conducted at 18.9°C and 21.1 °C; temperatures
at which the developmental rates of Wisconsin and Mexican eggs
differ. It would have proved interesting to run the hybridization
experiments at 19.8 °C, a temperature at which (interpolated from
figure 1) the rates of development are the same. An insufficiency
of frogs from Tamaulipas, Mexico frustrated efforts along this line,
but it appears likely that the same types and intensity of develop-
mental abnormalities would have been encountered if the reciprocal
crosses were conducted at 19-8 °C. The causative mechanism thus
remains problematic. Others have attributed heterogenic hybrid ab-
normalities to a variety of factors: altered activity or inactivation of
the sperm nucleus, disturbance of the nucleo-plasmic ratio, competition
of enzymes for limited substrates, etc.

Successful interpretation of our case and others is further com-
plicated by the demonstration that modifications in amphibian de-
velopment produced by direct action of a variety of chemical and
physical agents simulate in many ways inhibitions in hybrid develop-
ment (see Child, 1941).

No. 9 Volpe: Hybrid Inviability in Rana pipiens 123

Summary

1. The geographically extreme members of Rana pipiens from
Tamaulipas, Mexico, and Oshkosh, Wisconsin, are characterized by
different embryonic rates of development and embryonic temperature
tolerances, which may represent differential adaptations to the water
temperatures in nature to which the eggs of each group are subjected.

2. Experimental hybrids between Wisconsin and Mexican meadow
frogs show such marked embryonic abnormalities that none would
survive if the respective parental populations were to become sym-
patric and hybridize in nature. The evolutionary tenet that the
earliest indication of reproductive isolation appears as a by-product
of the genetic divergence of allopatric populations is supported.

3. The types of embryonic abnormalities in the reciprocal hybrids
follow definite patterns. Hybrid embryos derived from the cross
Mexican 2 X Wisconsin $ exhibit persistent blastocoeles, micro-
cephaly, fused suckers and olfactory pits, reduced mouthparts, and
atypical eyes. Hybrid embryos from the reverse cross, Wisconsin 2
X Mexican $ , possess enlarged yolk plugs, macrocephaly, over-
developed suckers, shortened bodies, and curved tails.

References Cited

Child, C. M. 1941. Patterns and Problems of Development. Univer-
sity of Chicago Press, Chicago, Illinois.

Koopman, K. F. 1950. Natural selection for reproductive isolation
between Drosophila pseudoobscura and Drosophila persimilis.
EvoL, 4: 135-148.

Moore, J. A. 1946. Incipient intraspecific isolating mechanisms in
Rana pipiens. Genetics, 31: 304-326.

1947. Hybridization between Rana pipiens from

Vermont and eastern Mexico. Proe. Nat. Acad. Sci., 33: 72-75.

1949a. Patterns of evolution in the genus Rana.

In: Jepsen, G. L., G. G. Simpson, and E. Mayr (eds.), Genetics,
Paleontology, and Evolution. Princeton University Press, Prince-
ton, New Jersey.

1949b. Geographic variation of adaptive char-
acters in Rana pipiens Schreber. EvoL, 3: 1-24.

Muller, H. J. 1942. Isolating mechanisms, evolution and temperature.
Biol. Symp., 6: 71-125.

Pollister, A. W. and J. A. Moore. 1937. Tables for the normal de-
velopment of Rana sylvatica. Anat. Rec, 68 : 489-496.

Volpe, E P. 1952. Physiological evidence for natural hybridization
of Bufo americanus and Bufo fowleri. EvoL, 6 : 393-406.

1953. Embryonic temperature adaptations and

relationships in toads. Physiol. ZooL, 26: 344-354.

flA- \\l<^»/ — V/e^-n^]

Jim ^®(DIL®OT

Volume 1, Number 10

April 30, 1954

THE BUTTERFLIES AND SKIPPERS OF LOUISIANA

EDWARD NELSON LAMBREMONT,

NEW ORLEANS.

HAY 1 1 1254

TULANE UNIVERSITY
NEW ORLEANS

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

William T. M. Forbes, Professor of Entomology, Cornell Uni-
versity, Ithaca, New York.

Austin H. Clark, Curator of Echinoderms, United States Na-
tional Museum, Washington, D. C.

Ralph L. Chermock, Associate Professor of Biology, University
of Alabama, University, Alabama.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
torial matters, and orders for individual numbers to the editor. Re-
mittances should be made payable to Tulane University.

When citing this series authors are requested to use the following
abbreviations: Tulane Stud. Zool.

Price for this number: $0.60.

Assistants to the Editor:
Carol L. Freret
Robert E. Gordon

George Henry Penn, Editor,
c/o Department of Zoology,
Tulane University,
New Orleans, U. S. A.

LIBRARY

WAY 1 1 1954

THE BUTTERFLIES AND SKIPPERS OF LOUISIANA1

EDWARD NELSON LAMBREMONT,
New Orleans

Prior to the summer of 1948 when a project to survey the butterflies
and skippers of Louisiana was started there existed only scattered
records and four parish lists. The oldest of these was published as a
catalog of the Lepidoptera of New Orleans and its vicinity by Ludwig
von Reizenstein in 1863. As originally published the records of von
Reizenstein could not have been included in this paper because the
basis of von Reizenstein's determination of species was unknown.
Twenty-six of the species, however, were checked against Morris
(1862) and are included with the other records. Six species, as yet
unidentifiable, are listed for general interest in Table 1.

Kopman (1903) published a popular account of the butterflies of
Louisiana in the New Orleans Times Democrat. Of the 24 species
recorded only three have definite localities noted and are included
herein. Hine (1904, 1906) listed the Lepidoptera of Cameron parish
in the southwestern corner of the state, including valid records for
15 species, and Montgomery (1932) listed 15 species from Madison
and Tensas parishes in the northeastern corner. Glick (1939) listed
eight species from Madison parish. All other publications deal with
only a few species.

From these publications a small, highly localized list may be com-
piled consisting of 43 species, including four type localities, in the
state.

The best and most recent local study was published since the initi-
ation of this survey. It consists of collections and notes made by
Jung (1950) within Orleans, Jefferson, Plaquemines, and St. Bernard
parishes. Thirty-seven species of butterflies and skippers are recorded,
17 of which had not been reported from the state previously.

Taking Jung's paper (1950) into consideration the state list num-
bered 60 species and subspecies. The present survey, concluded in
March 1951, adds 38 previously unrecorded species and subspecies,
bringing the total to 98 (Table 2). This is by no means to be
considered all the species that should be found, for many that are
known to occur in nearby areas have not been taken within Louisiana's
borders. The rich fauna of Texas and Mexico undoubtedly overlaps
part of Louisiana, particularly the southwest, and many of the Texas-
Mexican forms have been taken short distances from the Louisiana-
Texas line. In all probability the number of species would have
reached about 140 if the survey had been continued. Completion of
the survey would require considerable collecting in the state during
certain seasons of the year, particularly in the spring. On the basis

1 From a thesis submitted in partial fulfillment of the require-
ments for the degree of Master of Science of the Graduate School
of Tulane University.

128 Tulane Studies in Zoology Vol. 1

TABLE 1.

Species Recorded by von Reizenstein (1863) Which Were

Unidentifiable. Notes are Presented Exactly as They Appear

in the Original; Comments by the Writer are in Parentheses.

Goniopteryx Eclipsis.

Xanthidia Sylvatica, MSS Rare.

Heliconia diaphona Drur. Rare Western Lake Shore.
(Probably H. charitonius)

Euptoieta hegesia? Cram. City Park. On Passiflorae.
(Probably E. claudia)

Apatura myops, MSS. On elm trees.

Thecla io, MSS. City Park, rare. On camomilla.

of their overall distributional patterns approximately 40 additional
species may be anticipated to occur in Louisiana. These are listed
in Table 3, but otherwise are not mentioned in this paper.

Area Studied
Including islands in the Gulf of Mexico the total area of Louisiana
is 48,506 square miles, of which 3,097 square miles are water surface
(1,060 square miles of landlocked lakes). The entire state is within
Merriam's Austroriparian life zone; the northern portion within the
Carolinian area, and the southern portion in a semi-tropical region
called the Gulf Strip.

Viosca (1933) divided Louisiana into four main physiographic
areas which are illustrated in figure 1. Each of these supports a
characteristic flora. As will be noted later the distribution of certain
butterflies and skippers is correlated with these areas to some extent.
In addition to the natural vegetation there are many man-made com-
munities which tend to obscure the relation between natural vegeta-
tion and physiographic areas. Such man-made disturbance commun-
ities are: pine-hardwood, scrub-oak grassland, successional changes on
abandoned farmlands, and undeveloped agricultural land-reclamation
projects. These man-made disturbance areas support many small
flowering plants, both natural and horticultural, which are attractive
to butterflies and skippers.

With the exception of the coastal ridge and delta division all major
physiographic regions of Louisiana were sampled intensively during
the two-and-one-half year term of this survey. Those places within
short distances from New Orleans were concentrated upon, but three
major trips totalling more than 50 days in the field were taken to
collect in the northern uplands. The accompanying map (fig. 2)
shows the distribution of collections. To facilitate travel and in-
crease the range only a few areas were sampled over any length of
time. These include the Sabine parish area; Caddo and neighboring

No. 10

Lambremont: Butterflies and Skippers

129

Figure 1. Physiographic map of Louisiana from Viosca (1933). Re-
printed by permission of the copyright owner.

parishes; Lincoln-Jackson-Union parish area; Ouachita-Morehouse and
parishes east; Concordia parish and surrounding area; Allen parish
and all parishes east to the Mississippi river, excluding West Baton
Rouge; the Florida parishes; and New Orleans and vicinity.

Most of the collections were made during the summer months.
The only heavily collected sections during other times of the year
were the Jackson-Lincoln-Union area in the spring, Allen parish and
all parishes east to the Mississippi river in the fall, and the Florida
parishes and New Orleans and vicinity throughout the year.

The majority of the butterflies and skippers were taken by the
author from June 1948 to April 1951; records previous to this time
were contributed by other members of the Tulane University zoology
staff.

Systematic Treatment

Each species is considered individually. Each such treatment in-
cludes a summary of observations and Louisiana distribution records.
The arrangement of species follows Klots (1951).

Where specific data are required the records are listed alphabetically
by parish, each including the following: locality (within five miles

130

Tulane Studies in Zoology

Vol. 1

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No. 10

Lambremont: Butterflies and Skippers

133

Figure 2. Map showing distribution of localities from which mate-
rial was examined, or previously recorded; triangles = locali-
ties from the literature; dots = localities from the present
survey.

of nearest Post Office), date of collection, number and sex of speci-
mens, and institutional location of specimens in parentheses. Unless
otherwise specified all specimens are in the Tulane collection. Ab-
breviations of the institutions are: LPI = Louisiana Polytechnic In-
stitute, Ruston; LSM = Louisiana State Museum, New Orleans; SLI =
Southwestern Louisiana Institute, Lafayette; TU = Tulane University
("P" series are collections prior to 1947 with sparse data; lot numbers
P-955 through P-1101 consist of the James M. McArthur Collection).

Superfamily PAPILIONOIDEA, The Butterflies
Family Satyridae
Lethe portlandia portlandia (Fabricius), the Pearly Eye. —
The writer has not taken this species; however, Skinner (1897) de-
scribed a specimen captured in Louisiana as Ls creola. Dr. Austin H.
Clark informed the writer (in correspondence) that this supposed
creola is actually portlandia. Skinner's record is: St. Landry: Ope-
lousas, VII-3-1897.

Euptychia cymela cymela (Cramer). — The Little Wood-satyr

134 Tulane Studies in Zoology Vol. 1

is rather common in Louisiana during the spring and early summer;
earliest and latest dates recorded are April 1, 1950 (8 $ $ ) and June
24, 1950 (12). It is most often seen in woody places, particularly
in pinelands, and most of the specimens have been taken in the upland
pine areas of the state.

E. cymela cymela has not been recorded from Louisiana previously.

Twenty-four specimens (12 55,12 2 2) have been examined from

14 localities in the following parishes: Catahoula, Jackson, Lafayette

(SLI), LaSalle, Lincoln, Morehouse, Ouachita, St. Tammany, Union

and Washington.

Euptychia HERMES SOSYBius ( Fabricius ) . — The Carolinian Satyr
is undoubtedly the most abundant satyrid; it is probably rivaled by
only two other species, Eurema lisa and Phyciodes tharos, in being the
most common butterfly in Louisiana. This satyrid is particularly
abundant in woody areas, especially in pinelands and oak forests,
where it remains in quiet shaded spots rather than in the sunny hab-
itats of most other butterflies. This subspecies flies the year round;
specimens have been collected in all months except January and Au-
gust. Earliest and latest dates are February 10, 1943 (15) and De-
cember 25, 1949 (15).

One hundred and nine specimens (64 5 5, 45 2 2 ) have been
examined from 51 localities in the following parishes: Allen, As-
sumption, Catahoula, Claiborne, Evangeline, East Feliciana, Jackson,
Jefferson, Lafayette (SLI), Lincoln, Livingston, Morehouse, Orleans,
Ouachita, Pointe Coupee, Richland, Sabine, St. Helena, St. Landry, St.
Tammany, Tangipahoa, Union, Washington, West Carroll and West
Feliciana. Previously it had been recorded from Cameron (Hine,
1906) and Orleans parishes (von Reizenstein, 1863).

Euptychia areolata areolata (J. E. Smith), the Georgia
Satyr. — All specimens were collected in the longleaf pine flats north
of Lake Pontchartrain. Supposedly the species ranges as far west as
Texas, but the results of this survey indicate it must be rare west of the
Mississippi river, although it localizes and can be overlooked. It re-
sembles E. bermes in habits and flight by keeping close to the ground.

E. areolata areolata has not been recorded from Louisiana previous-
ly. The 31 specimens (15 5 5, 16 2 2), examined were collected
from the following localities: Livingston: Springville, IX-6-50 (TU
2184); St. Tammany: St. Tammany, V-12-49 (TU 1219); Hickory,
V-12-50, VI-2-50, VI-29-50, IX-30-50 (TU 1948, 1973, 2000, 2198);
Covington, IX-5-50, IX-8-50 (TU 2180, 2197); Folsom, IX-7-50
(TU 2187); Talisheek, IX-13-50 (TU 2200); Tangipahoa: Poncha-
toula, IX-6-50 (TU 2182); Robert, IX-6-50 (TU 2185); Washington:
Angie, VI-5-50 (TU 2034).

Euptychia gemma gemma (Hubner), the Jeweled Brown. —
This beautiful woodland butterfly resembles E. hermes closely in
habits, but it is considerably less common for only three specimens
(all females) have been taken in Louisiana. From the writer's expe-

No. 10 Lambremont: Butterflies and Skippers 135

rience the species occurs in pinelands, and several which eluded cap-
ture were seen in the longleaf pine flats of St. Tammany parish.

This species has not been recorded from Louisiana previously.
The three records are: Richland: Girard, VI-27-50 (TU 2081); St.
Tammany: Mandeville, X-26-49 (TU 1599); Tangipahoa: Tangi-
pahoa, IX-7-50 (TU 2192).

Cercyonis pegala pegala (Fabricius). — The Southern Wood-
nymph probably has a better protective coloration than any other
butterfly except its allied subspecies, C. pegala alope. The insect
usually may be found sitting on the sides of pine trees, and unless
disturbed it is often mistaken for a loose piece of bark. The un-
dersides of the wings are colored in such a way that they blend with
the surroundings so well that one must "beat" the trees with a stick
to flush them from hiding. When disturbed the butterfly quickly
flies a short distance to another tree, and the same procedure must
be repeated. To add to this collecting difficulty they are exception-
ally hard to capture in flight. Only a few have been collected in
Louisiana, perhaps because of their deceptiveness.

C. pegala pegala has not been recorded from Louisiana previously.
The three records are: St. Tammany: Covington, IX-8-50, 1 $ (TU
2197); Hickory, IX-13-50, 1 2 (TU 2198); Tangipahoa: Amite,
IX-7-50, 1 9 (TU 2195).

Cercyonis pegala alope (Fabricius), the Common Wood-
nymph. — In the past, authors have treated this subspecies as a sep-
arate species along with other forms of C. pegala. More recently
some have chosen to combine all as one large clinal species. Until
the pegala-alope complex is understood fully the writer prefers to
consider them as separate subspecies. The observations given for
typical pegala are much the same as those for alope.

C. pegala alope has not been recorded from Louisiana previously.
The only record is: LaSalle: Summerville, VI-12-50, 1 S (TU 2048).

Family Danaidae

Danaus plexippus plexippus (Linne). — The Monarch is one of
the most interesting butterflies of our fauna, and has attracted con-
siderable attention because of its migrations. For many years they
have been known to swarm (Saunders, 1871) and migrate.

The Louisiana collections show two annual peaks of abundance of
this subspecies: one in April, and the other during October and No-
vember. Judd (1950) reported a congregation of Monarchs on the
north shore of Lake Erie which occurred during the last four days of
August 1949, and Saverner (1908) reported swarms in Michigan
which lasted from September 4 through 15. Williams (1930)
quoted Shannon in saying that one of the four main routes of mi-
grating butterflies in North America is from Canada along Lake
Ontario and Lake Erie southward. Travelling at a speed of five to
seven miles per hour (Park, 1948), a swarm of monarchs leaving

136 Tulane Studies in Zoology Vol. 1

Canada in late August should reach the Gulf states in about a month
providing there are no high winds against them. This correlates
somewhat with our recorded peak of October-November, and also
with a large flight which the writer observed headed southward
October 10, 1949 near New Orleans, and one noted in November
1941 by Percy Viosca (newspaper article).

Originally this insect was more or less confined to North America,
but with time it has spread throughout the islands of the Atlantic,
and to most of the Pacific islands, including New Zealand and Aus-
tralia (Williams, 1930, 1937).

Forty-five specimens have been examined (27 $ $ , 1822) and
hundreds more have been observed in migratory flight. Collections
have been made throughout the year, except January and February;
earliest and latest dates are March 18, 1950 (15) and December
15, 1949 (1 $). The monarch has been recorded from 24 locali-
ties in the following parishes: Avoyelles (SLI), Franklin, Jefferson,
Lafayette (SLI), Lincoln, Livingston, Morehouse, Orleans, Ouachita,
Richland, St. Bernard, St. Charles, St. Helena, St. James, St. John the
Baptist, St. Landry, St. Tammany, Tangipahoa and West Feliciana.
Previously the monarch had been recorded from Cameron (Hine,
1906) and Orleans parishes (von Reizenstein, 1863; Shufeldt, 1884;
Jung, 1939, 1950).

Danaus plexippus melanippe (Hubner), the South American
Monarch. — This subspecies has caused much controversy in recent
years for the figure in Holland (1898; 1931 revised) given to rep-
resent D. plexippus plexippus is actually the subspecies melanippe.
Although melanippe is not typical for most of North America it has
been captured as an occasional vititor as far north as Illinois and New
York (Clark, 1941).

The Louisiana record is: Orleans: New Orleans, VIII-8-25, 1 $
(TU P-967). Previously it had been recorded from St. Tammany
parish by Clark (1941).

Danaus gilippus Berenice (Cramer). — The Queen is a rare
danaid in Louisiana. The few records show that this species flies
mostly during the warmer months. One female, however, was taken
in March, 1950, probably because the winter of that year was very
mild. Unlike the monarch this species is not migratory.

The six specimens examined were from the following localities:
Jefferson: Metairie, VIII-7-47 (TU 92); Bridgedale, VIII-25-50
(TU 2123); Orleans: New Orleans, XI-26-27, VII-1-32, 111-25-50
(TU P-970, P-982, 1825); SK Tammany: Mandeville, X-ll-49
(TU 1566). Previously it had been recorded from Cameron (Hine
1906) and Orleans parishes (Jung, 1939, 1950).

Family Nymphalidae
Heliconius charitonius tuckeri Comstock and Brown. —
The Zebra is not common in Louisiana, but it is seen occasionally
during the summer and early autumn in the southernmost parishes.

No. 10 Lambremont: Butterflies and Skippers 137

The few records from the state were obtained in August and October.
Although the writer has never seen this butterfly in the field, a few of
them were reported to him from the Mississippi delta region, the
southern extremity of Louisiana. Comstock and Brown (1950) re-
corded it from Texas. The larvae feed on passion flowers.

The few records are as fellows: Jefferson: Mississippi river bank
opposite New Orleans, X-18-1894 (Kopman, 1903); Orleans: New
Orleans (von Reizenstein, 1863); Terrebonne: Houma, VIII-9-19
(LSM).

Agraulis vanillae nigrior Michener, the Gulf Fritillary. —
This is the commonest fritillary in Louisiana. During the hot summer
months swarms of them may be seen hanging on blades of grass in
open fields. It flies from spring to early fall; earliest and latest dates
are March 22, 1950 and November 7, 1919.

The flight is exceptionally fast, and efforts at chasing this insect
on the wing usually prove unsuccessful. One trick it uses in elud-
ing capture is to fly up to about eight feet and suddenly reverse its
direction. It can change its course much faster than a running man,
and the writer has often found himself giving "chase" in the op-
posite direction from which the butterfly was traveling. However,
many specimens may be taken while they are resting. According to
Williams (1937) this butterfly is migratory.

The species apparently is more abundant in the southern parishes
of Louisiana. Forty-seven specimens (30 $ $, 17 2 2 ) have been
examined from 18 localities in the following parishes: Allen, Jeffer-
son, Lafayette (SLI), Livingston, Orleans, Plaquemines, Pointe Cou-
pee, Rapides, St. Bernard, St. Landry, St. Tammany and Terrebonne
(LSM). Previously it had been recorded from Cameron (Hine,
1906), Madison (Montgomery, 1932) and Orleans parishes (von
Reizenstein, 1863; Michener, 1947; Jung, 1950).

Dryas Julia delila (Fabricius), the Julia Butterfly. — McDun-
nough (1938) listed this species as of doubtful North American
occurrence, but Holland (1931) stated that it is an occasional visi-
tor in the extreme South. The genus is primarily a tropical one.

D. julia delila has not been recorded from Louisiana previously.
The one record is: Lafayette: Lafayette, XI-5-41, 1 $ (SLI).

Euptoieta Claudia Claudia (Cramer). — The Variegated Fritil-
lary is very common in Louisiana from early spring throughout the
summer; earliest and latest dates are March 31, 1950 (12) and Oc-
tober 16, 1949 (12). The larvae feed on passion flowers and
violets. Several years ago, in early July, the writer's attention was
called to a vacant lot in uptown New Orleans which was literally
"seeded" with chrysalids of this butterfly. Previous to the time that
this lot was infested there was a dense growth of passion flowers,
but after the feeding of thousands of caterpillars nothing remained
but bare twigs covered with chrysalids. Needless to say the emer-
gence of adults was sensational, and a good series of pupae was col-
lected and reared.

138 Tulane Studies in Zoology Vol. 1

Within Louisiana, E. claudia claudia appears to be more abundant
in the southern areas. Thirty-nine specimens {20$$, 19 5?)
have been examined from 15 localities in the following parishes:
Assumption, Concordia, Evangeline, Jefferson, Lafayette (SLI), La-
fourche, Orleans, Ouachita, St. Bernard, St. Landry (SLI) and St.
Martin (SLI). Previously it had been recorded from Cameron
(Hine, 1906), Madison (Montgomery, 1932) and Orleans parishes
(von Reizenstein, 1863; Jung, 1950).

Speyeria cybele (Fabricius), the Great Spangled Fritillary. —
The single Louisiana specimen was taken in the coastal area in early
October, and as the species is not common to the state, perhaps it
was a straggler southward during the cooler months.

S. cybele has not been recorded from the state previously. The
one record is: Lafayette: Lafayette, X-2-31, 1 $ (SLI).

Melitaea nycteis nycteis Doubleday, the Silvery Checkerspot.
— Holland (1931) stated that M* nycteis ranges from Maine to western
North Carolina, and westward to the Rocky Mountains. He did not
say whether it comes below the latitude of North Carolina, or stays
north of there. Klots (1951) also did not mention its occurrence
south of the mountains of North Carolina, Missouri and Kansas. The
Louisiana records may be considered southern range extensions, but
since the species is not abundant in the state it appears that Louisiana
is at the southern tip of its range. A single specimen was collected
in the Sabine river area (near Texas), and another near the Pearl
river (near Mississippi), at opposite sides of the state. Three speci-
mens were seen at the latter location, but only one was taken. The
flight is fast and erratic.

Ms nycteis nycteis has not been recorded from Louisiana previously.
The two records are: Sabine: Zwolle, VIII-2-49, 1 5 (TU 1381);
Washington: Richardson, IX-7-50, 1 $ (TU 2188).

Phyciodes gorgone (Hubner), the Gorgone Crescent-spot. —
This crescent-spot, though closely allied to it, cannot begin to ap-
proach P. tharos in abundance. It flies from spring to early fall in
sunny places, but does not seem to be restricted to any part of the
state. Earliest and latest dates are March 7, 1951 (1 $ ) and Oc-
tober 16, 1949 (1 $).

Twenty-three specimens (14 $ $ , 9% 9) have been examined
from 14 localities in the following parishes: Allen, Assumption, Bos-
sier, DeSoto, Evangeline, Natchitoches, Pointe Coupee, Rapides, St.
Charles, St. Tammany and Union. Previously it had been recorded
from Cameron (Hine, 1906) and Orleans parishes (Jung, 1950).

Phyciodes tharos tharos (Drury). — The Pearly Crescent-spot
is by far the most abundant butterfly in Louisiana. With the excep-
tion of the coldest days it flies throughout the year (it has been col-
lected in all twelve months), not only in bright open fields, but in
densely shaded forests as well, and is known to cluster around wet
spots on the ground (Scudder, 1889). When at rest this butterfly

No. 10 Lambremont: Butterflies and Skippers 139

has the peculiar habit of slowly raising and lowering its wings. The
larvae feed on plants of the family Compositae, particularly the asters.
One hundred and thirty-one specimens (74 $ $ , 57 2 2 ) have
been examined from 68 localities in the following parishes: Allen,
Assumption, Catahoula, Claiborne, Concordia, DeSoto, East Baton
Rouge, East Feliciana, Evangeline, Jackson, Jefferson, Lafayette (SLI),
Lafourche, LaSalle, Lincoln, Livingston, Morehouse, Natchitoches,
Orleans, Ouachita, Pointe Coupee, Rapides, Richland, Sabine, St.
Charles, St. Helena, St. James, St. Landry, St. Tammany, Tangipahoa,
Union, Washington, Webster and West Carroll. Previously it had
been recorded from Madison (Montgomery, 1932) and Orleans
parishes (Jung, 1950).

Polygonia interrogations (Fabricius), the Question Mark. —
This species is exceptionally nervous and fast in flight, which per-
haps accounts for the scarcity of records. In the writer's opinion
each record obtained equals at least two or three uncaught butter-
flies, and from sight-records it appears that this species is rather
abundant in Louisiana during the spring and summer months. Ear-
liest and latest dates are March 10, 1949 (1 ?) and October 15,
1926 (1 2).

This butterfly frequents open spots and forest edges. Scudder
(1889) notes that it often sucks the sap flowing from wounded trees
and decaying fruits, and the writer has taken adult specimens feed-
ing on decaying fish and turtle meat. The larvae feed on a great
variety of plants, particularly of the family Urticaceae.

Eighteen specimens (7##, 1199) have been examined from
eight localities in the following parishes: Jefferson, Lafayette (SLI),
Orleans, Ouachita, Sabine, Terrebonne (LSM), Washington and
West Carroll. Previously it had been recorded from Orleans par-
ish by Jung (1950).

Nymphalis ANTIOPA ANTIOPA (Linne), the Mourning Cloak. —
This beautiful butterfly is primarily a species of the northern states,
appearing only occasionally in Louisiana during the late summer and
fall months. It has not been recorded from the state previously.

The three records are: Lincoln: Ruston (LPI); Orleans: New
Orleans, IX-25-31, 1 9 (TU P-977), and VIII-25-32, 1 2 (TU
P-983).

Vanessa atalanta atalanta (Linne), the Red Admiral. — This
species is well-named atalanta for it is one of the speediest of the
nymphalids. All of the specimens were caught while occupied with
feeding. The larvae feed on nettles (Urtica spp.)

In Louisiana, V. atalanta atalanta is never too abundant, but it
may be seen almost throughout the year, although more often dur-
ing the warmer months. Earliest and latest dates are January 30,
1950 (3^,299) and October 28, 1928 (1 2). Jung (1950)
stated that he found it to be abundant in drained swamp areas near
New Orleans.

140 Tulane Studies in Zoology Vol. 1

Twenty-one specimens (8 $ $ , 13 2 5) have been examined
from eight localities in the following parishes: Claiborne, Jefferson,
Lafayette (SLI), Orleans, Ouachita, St. Tammany, Webster and West
Feliciana. It had been reported previously from Orleans parish by
Jung (1939, 1950).

Vanessa cardui CARDUI (Linne), the Painted Lady. — In Louisi-
ana this butterfly is not quite as abundant as the related species at-
alanta and virginiensis, the few records indicating that it flies from
spring throughout the summer and autumn. Earliest and latest dates
are May 3, 1924 (15) and October 18, 1949 (1 $ ).

Williams (1937) described the migrations of cardui, noting that
in 1931 their abundance in the northern states brought help to the
farmers because of the vast destruction of thistles, the food plant of
their larvae. Yet in other years they do considerable damage to
artichokes.

The specimens taken in the state were from open, sunny fields, and
on the shrubs bordering forests. Only one was taken on thistle.

The localities are: Caddo: Shreveport, VIII-5-49, 1 5 (TU 1432);
Orleans: New Orleans, V-3-24, 1 5 (TU P-957); St. Bernard: Shell
Beach, X-18-49, 1 $ (TU 1595); St. Landry: Cortableu, X-16-49,
1 5 (TU 1580). Previously it had been recorded from Cameron
(Hine, 1906) and Orleans parishes (von Reizenstein, 1863).

Vanessa virginiensis (Drury). — Hunter's Butterfly is fairly
common in Louisiana through most of the year; earliest and latest
dates are January 21, 1950 (3 $ $) and October 10, 1930 (15).
Most of the specimens were taken while feeding, for like its ally, V.
atalanta, this species is a strong flier. The larvae feed on various
species of the family Compositae.

Nineteen specimens (11 $ S , 8 5 5) have been examined from
nine localities in the following parishes: Catahoula, East Feliciana,
Jackson, Lafayette (SLI), Orleans, St. Charles, St. Tammany, Wash-
ington and West Feliciana. Previously it had been recorded from
Madison (Montgomery, 1932) and Orleans parishes (von Reizen-
stein, 1863; Jung, 1939, 1950).

Precis evarete coenia (Hubner), the Buckeye. — This common
butterfly of the southern states probably is familiar to everyone. In
Louisiana it is very abundant throughout the spring, summer and
autumn; earliest and latest dates are January 21, 1950 (1 $ ) and No-
vember 8, 1949 (15,15). Of all that were collected in the state
only a few were obtained while they were flying or feeding, for the
insect seems to prefer to sit on hard, dry mud in full sunlight, slowly
moving its wings. During a recent visit to Lacombe, La. the writer
saw hundreds of buckeyes sitting on a large mound of dry, yellow
clay.

Holland (1931) stated that the larvae in the later stages feed on
plantains and Scudder (1889) reported them on Gerardia purpurea
and on other species of the family Scrophulariaceae.

No. 10 Lambremont: Butterflies and Skippers 141

Forty-seven specimens (31 ^ ^ , 1692) have been examined
from 31 localities in the following parishes: Allen, Claiborne, East
Baton Rouge, Evangeline, Jefferson, Lafayette (SLI), Livingston,
Natchitoches, Orleans, Ouachita, Rapides, St. Bernard, St. Charles,
St. James, St. John the Baptist, St. Landry, St. Tammany, Tangipahoa,
Union and Washington. Previously it had been recorded from
Madison (Glick, 1939) and Orleans parishes (von Reizenstein, 1863;
Jung, 1950).

Limenitis ARTHEMis ASTYANAX (Fabricius), the Red-spotted Pur-
ple.— This butterfly is very fast in flight, and if disturbed will soar
into the air. Most of the specimens were obtained in damp, swampy
places where they often sit on soft mud, or upon the leaves of shrubs
and small trees. The larvae feed on cherry, apple, hackberry, willow
and related shrubs and trees (Holland, 1931).

The species flies throughout the summer and early fall; earliest and
latest dates are May 31, 1949 (15) and October 15, 1949 (15).

Thirteen specimens (6 $ $ , 7 5?) have been examined from
nine localities in the following parishes: Catahoula, Claiborne, East
Feliciana, Livingston, Orleans, Tangipahoa, Terrebonne (LSM) and
West Feliciana. Previously it had been recorded from St. Tammany
parish by Jung (1950).

Limenitis archippus watsoni (Dos Passos), the Viceroy. —
This butterfly exhibits one of the best known examples of protective
mimicry. It looks almost exactly like the monarch, Danaus plexippus,
a species protected from insectivores by the unpleasant taste it pos-
sesses from feeding on milkweeds. An interesting fact concerning
the viceroy is that throughout its range are found subspecies each of
which mimics the prevalent danaid of the area (Chermock, 1947).

L. archippus ranges throughout North America into the Gulf States
where the subspecies floridensis, in the east, and watsoni, in the west,
occur. The latter was described by Dos Passos (1938) from speci-
mens caught at Alexandria, La. In the state this subspecies is very
common during the spring, summer and autumn months; earliest
and latest dates are April 24, 1948 (15) and November 12, 1949
(1 5).

The flight, usually within five feet of the ground, is fast, but lei-
surely, and the butterfly is not too difficult to capture as it flies, or
when it feeds on flowers. The larvae feed on various species of wil-
lows, although occasionally they feed on plum and cherry (Scudder,
1889).

Thirty-three specimens (2455,955), have been examined
from 22 localities in the following parishes: Allen, Assumption, Clai-
borne, Evangeline, East Baton Rouge, Jefferson, Lafayette (SLI),
Lafourche, Livingston, Orleans, Pointe Coupee, St. Bernard, St.
Charles, St. Helena, St. James, St. Landry and Tangipahoa. Pre-
viously it had been recorded from Orleans (Jung, 1950), Rapides
(Dos Passos, 1938) and Tensas parishes (Montgomery, 1932).

142 Tulane Studies in Zoology Vol. 1

Anaea andria (Scudder). — The Goatweed Butterfly is very com-
mon in Louisiana during the spring and early summer, but because
of its rapid flight and protective coloration it is difficult to capture.
Earliest and latest dates are February 2, 1929 (12) and October
20, 1939 (1 $). When resting this butterfly usually sits on the
ground among dead leaves showing the under surfaces of the wings
which blend well with the surroundings. Several individuals have
been taken on the trunks of sycamore trees; others undoubtedly blend-
ed with the loose bark and were not seen. The caterpillars feed on
the goatweed (Croton capitatum) .

As andria appears to be slightly more abundant in northern Louisi-
ana. Most of the specimens were taken in the hardwood areas. Eight-
een specimens (9 S S , 9 9 2 ) were examined from 13 localities in
the following parishes: Catahoula, Claiborne, Concordia, Evangeline
(SLI), Lafayette (SLI), Morehouse, Orleans, Pointe Coupee, Winn
and Union. Previously it had been recorded from Cameron (Hine,
1906), Madison (Montgomery, 1932), Orleans (Jung, 1950) and
Rapides parishes (Johnson and Comstock, 1941).

ASTEROCAMPA CLYTON CLYTON (BOISDUVAL AND LeCONTE). —
The Tawny Emperor is not very common in Louisiana. The writer has
seen two females, and von Reizenstein (1863) and Jung (1950) re-
ported it from New Orleans. The butterfly is very fast in flight and
is difficult to catch. The larvae feed on leaves of the hackberry (Cel-
tis mississippiensis ) .

The Louisiana records are: Lafayette: Lafayette, VII-20-41, 1 9
(SLI); Ouachita: Sterlington, VI-23-50, 1 $ (TU 2072).

ASTEROCAMPA CELTIS CELTIS (BOISDUVAL AND LeCONTE). — In

Louisiana the Hackberry Butterfly is very common during the hot sum-
mer months, but due to the difficulty of capturing it only a few have
been taken. Earliest and latest dates are June 8, 1947 (1 $) and Au-
gust 3, 1931 (15). The flight is exceptionally rapid, high around
the tops of bushes and trees. The larval food is hackberry.

Twenty-one specimens (9 $ $ , 12 2 2 ) have been examined
from seven localities in the following parishes: Concordia, Lafa-
yette (SLI), Ouachita, Richland, and Sabine. Previously it had been
recorded from Orleans parish by von Reizenstein (1863) and Jung
(1950).

ASTEROCAMPA CELTIS ALICIA (Edwards). — Apparently in Louisi-
ana this subspecies of the hackberry butterfly is quite restricted in its
range, whereas A. celtis celtis is the dominant subspecies throughout
the state.

The writer saw many of these butterflies darting among the
branches of hackberry trees along the Mississippi river batture near
New Orleans. On that occasion two males were captured, but not
without considerable effort. The three specimens examined were
all taken in Orleans parish as follows: VII-10-32, 1 $ (TU P-982)
and IX-1-48, 2 $ $ (TU 1258).

No. 10 Lambremont: Butterflies and Skippers 143

Family LlBYTHEiDAE

LlBYTHEANA BACHMANNII BACHMANNII (KlRTLAND), the Snout

Butterfly. — This species is very abundant in Louisiana during the hot
summer months, but appears to fly at least occasionally even during
the winter as the writer has observed a few individuals in December
and January. Other than these "off season" observations the earliest
and latest dates are May 19, 1950 (1 $ ) and August 27, 1949 (H).
They may be found in swarms along roadside pools of water and damp
places on the ground. Williams (1937) described a southeasterly
migration of these "snouts" in September, which extended over a
front of 250 miles. He estimated that one and a quarter million
butterflies passed every minute on the whole front. The larvae feed
on hackberry trees.

Twenty-four specimens (17 5 5,7 22 ) have been examined from
14 localities in the following parishes: Catahoula, Concordia, East
Feliciana, Lafayette (SLI), Morehouse, Orleans, Sabine, St. Bernard,
Vermilion (SLI), Washington and West Feliciana. Previously it
had been recorded from Cameron (Hine, 1906), Madison (Mont-
gomery, 1932) and Orleans parishes (Jung, 1939, 1950).

Family Riodinidae

Calephelis virginienis (Guerin), the Little Metal-mark. — The
writer has captured a single male of this tiny butterfly from the
Florida parish area, in the easternmost section of the state. Little is
known of the immature stages of this species.

The lone record is: St. Tammany: St. Tammany, 111-29-51, 1 5
(TU 2349). Previously it had been recorded from Opelousas in St.
Landry parish by Holland (1931) under the synonym C. louisiana.

Family Lycaenidae

Atlides halesus halesus (Cramer), the Great Purple Hair-streak.

— The writer has not taken this beautiful insect, but Jung (1950)

reported it from New Orleans. The one specimen examined was

also collected in New Orleans, November 11, 1933, 1 5 (TU P-991).

Strymon CECROPS (Fabricius). — The Cecrops Hair-streak is
common in Louisiana during the spring and summer months, and is
easily taken while it feeds on flowers. Earliest and latest dates are
January 30, 1950 (15) and October 16, 1949 (H). A habit
peculiar to the lycaenids consisting of "cleaning" or rubbing together
the upper surfaces of the wings while at rest, is practiced strongly
by this species.

Twelve specimens (1055,2 29) have been examined from
twelve localities in the following parishes: Allen, East Feliciana,
Franklin, Lafayette (SLI), Madison, Orleans, Ouachita, Pointe Cou-
pee, Sabine, St. Tammany and Tangipahoa. Previously it had been
recorded from Caddo (Field, 1941) and Madison parishes (Mont-
gomery, 1932).

Strymon melinus melinus (Hubner), the Common Hair-streak.
— This species is very common in Louisiana. The butterfly is very

144 Tulane Studies in Zoology Vol- 1

swift in flight but is taken easily while feeding on wild flowers. It
flies throughout the spring, summer and early fall; earliest and latest
dates are March 10, 1943 (1 $ ) and October 14, 1949 (12). How-
ard (1895) described the damage to bean plants caused by the
larvae.

Twenty-three specimens (15 3 #,8 9 5) have been examined
from 16 localities in the following parishes: Catahoula, Evangeline,
Jefferson, Lafayette (SLI), Lincoln, Livingston, Natchitoches, Or-
leans, St. Bernard, St. Tammany, Tangipahoa and Washington. Pre-
viously it had been recorded from Cameron (Hine, 1906) and Or-
leans parishes (Jung, 1950).

Strymon ONTARIO AUTOLYCUS (Edwards). — The Northern Hair-
streak has been taken only once in Louisiana. Klots (1951) stated
that it is not common, but may be caught easily while it feeds. The
larva feeds on oak.

S. Ontario autolycus has not been recorded from Louisiana previously.
The Louisiana record is: Orleans: New Orleans, V-5-51, 1 2 (TU
3010).

Strymon liparops liparops (Boisduval and LeConte), the
Striped Hair-streak.— Scudder (1889) stated that this butterfly is
usually localized near thickets and swamps. The only specimen
from Louisiana was taken in a dense upland hardwood forest with
a very thick undergrowth of vines.

S. liparops liparops has not been recorded from Louisiana pre-
viously. The lone record is: LaSalle: Summerville, VI- 12-50, (TU
2049),

Incisalia irus (Godart), the Hoary Elfin. — This species is
included on the basis of Skinner's report (1907), of it from the
state, although the record is rather dubious. Holland (1931) stated
that the species is always rather rare, but has a wide east-west distri-
bution in the latitude of New England. This means that Skinner
either published an exceptional range extension, or that he was con-
fused with other nearly related species; most likely his specimen was
Z„ henrici, with which irus has often been confused.

Feniseca tarquinius tarquinius (Fabricius), the Harvester. —
The writer has taken only one specimen of this species, a female
which was captured in a dense hardwood forest on the banks of the
Ouachita river in northeastern Louisiana. The butterfly was flying
about four feet from the ground in a very nervous manner and was
taken with difficulty. The larvae of this butterfly are of some eco-
nomic importance because they feed on woolly aphids, particularly
those of the alder and beech.

The single record is: Richland: 7 miles north of Girard, VI-27-50,

1 $ (TU 2084). Previously the species was recorded from Caddo

(Scudder, 1889), Orleans (von Reizenstein, 1863) and Webster

parishes (Johnson, 1886).

Hemiargus isolus (Reakirt), Reakirt's Blue. — Klots (1951)

No. 10 Lambremont: Butterflies and Skippers 145

gives the range as "Mexico, north to Nebraska, almost entirely west
of the Mississippi river." The few specimens collected in Louisiana
certainly appear to be at the eastern edge of the species' range, and
may be considered an extension as far as the southern portion of the
range is concerned.

H. isolus has not been recorded from Louisiana previously. The
records are: Concordia: 13 miles northwest of Ferriday, VI-16-50,
1 $ (TU 2056); Morehouse: 12 miles east of Mer Rouge, VI-19-50,
3 S $, 1 5 (TU 2063).

Everes COMYNTAS COMYNTAS (Godart). — The Eastern Tailed-blue
is very abundant in Louisiana during the spring and summer months;
earliest and latest dates are January 2, 1950 ( 1 2 ) and October 14,
1949 ( 1 5 ) . The writer has seen most specimens along the edges
of forests in partly shaded areas and on the ground around moist
places. Their quick flight and small size make them hard to catch.

Sixty-five specimens (38 $ $ , 27 2 $ ) have been examined from
25 localities in the following parishes: Catahoula, Evangeline, East
Feliciana, Lafayette (SLI), LaSalle, Lincoln, Madison, Morehouse,
Natchitoches, Ouachita, Richland, Sabine, St. Charles, St. Tammany,
Union, Washington and West Carroll. Previously it had been re-
corded from Madison parish by Montgomery (1932).

Celastrina argiolus pseudargiolus (Boisduval and Le-
CONTE). — The Spring Azure apparently is rare in Louisiana. The
few specimens collected were taken in the longleaf pine flats of the
Florida parishes. Scudder ( 1889) stated that the range does not touch
the Gulf of Mexico; hence this species may not be found in the coastal
parishes.

C. argiolus pseudargiolus has not been recorded from Louisiana
previously. The two records are East Feliciana: Felixville, VI-8-50,
3 2 $ (TU 2037); Washington: Angie, VI-4-50, 1 $ (TU 2032).

Family Papilionidae

Papilio polyxenes asterias (Stoll). — The Common American
Swallowtail apparently is not confined to any particular part of Lou-
isiana. It is rather abundant throughout the spring and summer;
earliest and latest dates are January 1, 1950 (15) and September 7,
1926 (12). It is a frequent visitor to flowers; specimens have been
taken on roses, clover and numerous umbelliferous plants. It has
some economic importance as the caterpillars may feed on celery,
dill, parsnip, carrots, caraway and parsley.

Twenty- three specimens (14 5 5,9?$) have been examined
from ten localities in the following parishes: Claiborne, Franklin,
Jefferson, Lafayette (SLI), Orleans, Richland, St. Charles, Tensas, Ver-
milion (SLI) and Washington. Previously it had been recorded
from Madison (Montgomery, 1932) and Orleans parishes (von
Reizenstein, 1863; Jung, 1939, 1950.

Papilio cresphontes cresphontes (Cramer). — The Giant Swal-
lowtail, one of largest of the North American butterflies, is of eco-

146 Tulane Studies in Zoology Vol. 1

nomic importance to fruit growers for the caterpillar, the "Orange-
puppy," feeds on citrus trees. In Louisiana it is more abundant in
the southernmost areas. Rothschild and Jordan (1906) record the
species from "Louisiana."

Saverner (1908) reported badly damaged P. cresphontes and P.
troilus migrating southward along with monarchs during August; this
may have been the situation of two cresphontes captured in north
Louisiana during August which were accompanying a southerly mi-
gration of monarchs. In Louisiana the species flies from spring to
early fall; earliest and latest dates are March 2, 1949 (1 $ , 1 ? ) and
November 13, 1911 (1 5).

Twenty-six specimens, (13 S $ , 13 5 $) have been examined from
eleven localities in the following parishes: Claiborne, Jefferson, Laf-
ayette (SLI), Livingston, Orleans, Plaquemines, St. Tammany, Ter-
rebonne (LSM) and Vermilion (SLI). Previously it had been re-
corded from Cameron (Hine, 1906) Orleans (von Reizenstein, 1863;
Jung, 1939, 1950) Plaquemines (Jung, 1950), St. Bernard (Jung,
1950) and Terrebonne parishes (Comstock, 1882).

Papilio GLAUCUS GLAUCUS (Linne). — The Tiger Swallowtail dis-
plays strong sexual dimorphism. In the southern part of its range
some of the females are dark brown rather than black and yellow.
According to Klots (1951) this melanism occurs in about 50 percent
of the females, but he states that the brown form is less common or
nonexistent in the extreme south. All of the females that the writer
has seen from Louisiana show this dark brown coloration. Some of
the males are larger and tend to show the darker yellow color of
P. glaucus australis, although none of the specimens appear clearly to
be that subspecies.

The species is a strong flier and is difficult to catch in flight;
however, the writer found them easy to collect while they were feed-
ing on the thistle (Cirsium horridulum). Riley and Howard (1895)
reported the caterpillars feeding on the leaves of camphor trees in
Louisiana.

The species has been collected in all months of the year except
January and December; earliest and latest dates for adults are Febru-
ary 18, 1950 (!&$)_ and November 16, 1939 (15).

Twenty-three specimens (YJ $ $ , 6 5 9) have been examined
from twelve localities in the following parishes: Assumption, Bossier,
Claiborne, Lafayette (SLI), Madison, Morehouse, Natchitoches, Or-
leans, Sabine, St. Charles, St. Tammany and Washington. Previously
it had been recorded from Jefferson (Jung, 1950), Morehouse (Riley
and Howard, 1895), Orleans (von Reizenstein, 1863) and St. Tam-
many parishes (Jung, 1950).

Papilio troilus troilus (Linne). — The Spicebush Swallowtail
is the most common member of its genus in Louisiana. It flies from
early spring to late summer; earliest and latest dates are February
23, 1949 (3 $ $) and September 16, 1945 (1$). The larvae and
adults feed on spicebush, sassafras and other Lauraceae.

No. 10 Lambremont: Butterflies and Skippers 147

Saverner (1908) stated that P. troilus was seen migrating south-
ward with the monarch (Danaus plexippus) in August, but no evi-
dence of migrations of this swallowtail have been noted as yet in
Louisiana.

Forty-one specimens (27 $ $ , 14 9 9 ) have been examined from
eleven localities in the following parishes: Caddo, DeSoto, Jackson,
Jefferson, Lafayette (SLI), LaSalle, Orleans, Ouachita, Sabine, St.
Tammany and Union. Previously it had been recorded only from
Orleans parish by von Reizenstein (1863) and Jung (1939, 1950).

Papilio troilus ilioneus J. E. Smith. — While many of the
spicebush swallowtails from Louisiana are more typical of the sub-
species troilus, three specimens possess very definite characteristics of
subspecies ilioneus. However, the majority collected have character-
istics common to either or both of the subspecies, due to the clinal
nature of the species.

P. troilus ilioneus has not been recorded from Louisiana previously.
The records are: Orleans: New Orleans, III-4-49, 2 9 9 (TU 1110);
Ouachita: Sterlington, VI-23-50, 1 $ (TU 2072).

Papilio palamedes palamedes (Drury). — The Palamedes Swal-
lowtail is a rather rare insect in Louisiana, records of this species
being confined to the coastal parishes. As this butterfly is a strong
flier and difficult to capture, adults are more readily obtained by
rearing them from the larval stage. The few specimens collected
were taken in the months of April, June, July, August and October;
earliest and latest dates are April 1, 1918 and October 13, 1934.

Nine specimens (5 $ $ , A 9 9) were examined from three local-
ities in the following parishes: Lafayette (SLI), Orleans and Terre-
bonne (LSM). Previously it had been recorded from Orleans parish
by von Reizenstein (1863), Rothschild and Jordan (1906) and
Jung (1950).

Graphium marcellus marcellus (Cramer), the Pawpaw
Butterfly. — Although this species is fairly common in Louisiana dur-
ing the early summer months it is exceedingly difficult to take with
a net. A few specimens were obtained, however, after many others
had eluded capture. Earliest and latest dates were April 1, 1950
(1 $ ) and September 10, 1947 (1 9). Late in the survey it was
learned that this butterfly usually picks an orbit to follow in its
flight, and if the collector misses it on the first try he has only to
wait a few minutes for the same individual to return, flying almost
exactly the same course.

Within Louisiana marcellus apparently is rare in the coastal and
Florida parish areas, and all of the specimens were obtained from
the vicinity of the Mississippi river westward and northward. It is
more abundant in the upland areas of the state. Its scarcity in
southern Louisiana may be due to the lack of its food plant, the
pawpaw (Asimine triloba and A. parviflora) , in the coastal region.

Eleven specimens (6 $ $ , 5 9 9) were examined from five local-

148 Tulane Studies in Zoology Vol. 1

ities in the following parishes: Catahoula, Lafayette (SLI), Sabine,
Union and West Feliciana. Previously it had been recorded from
Orleans parish by Kopman (1903) under the name Papilio ajax.

BATTUS PHILENOR philenor (Linne). — The Pipe-vine Swallow-
tail is most common during the summer months in Louisiana, but
it has a long flying season; earliest and latest dates are February 1,
1949 (1 $) and October 16, 1940 (15). Holland (1931) states
that philenor has been observed ovipositing as late as October. The
species is exceptionally fast in flight, perhaps accounting for the
scarcity of records. The larvae generally feed on Dutchman's pipe
or pipe-vine (Aristolochia serpentaria and A. durior).

Seventeen specimens (13 $ $ , 4 2 9) have been examined from
eleven localities in the following parishes: Bossier, Lafayette (SLI),
Morehouse, Ouachita, Rapides, Sabine, St. Tammany, Union and
Washington. Previously it had been recorded from Cameron (Hine,
1906), Orleans (von Reizenstein, 1863), St. John the Baptist (Jung,
1950), St. Mary (Kopman, 1903), and St. Tammany parishes (Jung,
1950).

Battus polydamus lucayus (Linne), the Polydamus Butter-
fly.— The writer has not seen this butterfly in Louisiana, but von
Reizenstein (1863) reported it from New Orleans. According to
his observations it was very rare, usually found feeding on Virginia
snake root and Dutchman's pipe. McDunnough (1938) doubted
its occurrence in North America, but Klots (1951) stated that it
occurs in the Gulf states from Florida to Texas.

Family Pieridae

Anthocharis genutia genutia (Fabricius), the Falcate Orange-
tip. — In Louisiana this species apparently is confined to the upland
regions of the northern part of the State. Scudder (1889) stated that
the butterfly has never been very abundant. From the writer's experi-
ence it seems that this species is a very weak flier, and is easy to
collect, although Scudder (1889) reported otherwise.

A. genutia has not been recorded from Louisiana previously. The
three records are: Jackson: Kelleys, IV-1-50, 1 $ (TU 1846);
Chatham, IV-1-50, 2 $ $ (TU 1849); Lincoln: Downsville, IV-1-50,
15,1? (TU 1859).

Colias eurytheme eurytheme Boisduval. — The Eurytheme
Sulphur can be found in all parts of Louisiana, but it is not as abundant
in the coastal area as it is in the pinelands and uplands, especially in
open grass or clover-covered fields. This species flies throughout most
of the year; earliest and latest dates are January 21, 1950 (2 8$) and
November 8, 1939 (1 2 ). The larvae feed occasionally on Trifolium
repens and other related species of this genus, but Scudder (1889)
reported that the caterpillars would not eat red clover. The favorite
food, however, is the introduced alfalfa (Medicago sp.), on which it
is a serious pest.

With the exception of C. a. pseudargiolus, a lycaenid, C. eurytheme

No. 10 Lambremont: Butterflies and Skippers 149

is probably the most variable species in our butterfly fauna. The
literature is filled with descriptions of the many sexual and seasonal
forms. These have been considered at times to be distinct species,
and as such have presented a very puzzling problem. However, in
recent years much has been done by Hovanitz (1943, 1944) to study
the Colias complex; hybridization and female albinism have been
studied also by Gerould (1946).

Sixty specimens (26 $ $ , 34 2 2 ) have been examined from 22
localities in the following parishes: Assumption, Claiborne, Concordia,
East Baton Rouge, Evangeline, Lafayette (SLI), Morehouse, Natchi-
toches, Orleans, Ouachita, Rapides, Richland, and St. Charles. Pre-
viously it had been recorded from East Baton Rouge (Floyd, 1941),
Madison (Montgomery, 1932; Glick, 1939) and Orleans parishes
(Scudder, 1889; Jung, 1939, 1950).

Colias philodice philodice (Latreille), the Common Sul-
phur.— The common name of this butterfly implies abundance, but it
is rather rare in Louisiana, and although it has a wide range through-
out North America one specimen has thus far been taken in the
state. From what can be discerned from Scudder (1889) it is not
numerous at the southern extremities of its range.

C. philodice philodice has not been recorded from Louisiana pre-
viously. The lone record is: Lafayette: Lafayette, XI-30-36, 1 $
(SLI).

Zerene cesonia (Stoll), the Southern Dog-face. — The writer
has examined only one specimen from Louisiana, and it has been
recorded by von Reizenstein (1863) from New Orleans. The fact
that this butterfly flies very fast and often high off the ground may
account for the scarcity of records. The species frequents pinelands,
and has been seen by the writer in the pine flats of the Florida parishes
in August, but on that occasion the individuals were flying fast and
high above the ground, and none were caught.

The single record is: Lafayette: Lafayette, XI-12-28, 1 $ (SLI).

Phoebis sennae eubule (Linne). — The Cloudless Sulphur is
very abundant during most of the year throughout Louisiana. Earliest
and latest dates are March 4, 1934 (1 2 ) and November 1, 1949
( 1 $ ) . It is an exceptionally strong flier, and is migratory. Great
swarms of them have been observed far out at sea, flying just above
the waves (Williams, 1937). During courtship when the male and
female "pinwheel" high above the ground they are especially hard
to capture, yet when feeding they are so occupied that the writer has
taken many by hand. The larvae feed on Cassia spp.

Forty-six specimens (25 $ $ , 21 22) have been examined from
32 localities in the following parishes: Allen, Assumption, Bossier,
Claiborne, Concordia, East Baton Rouge, Evangeline, Franklin, Jeffer-
son, Lafayette (SLI), Orleans, Ouachita, Pointe Coupee, Rapides,
Red River, Richland, Sabine, St. Bernard, St. Helena, St. Tammany,
Tangipahoa (SLI, TU) and Washington. Previously it had been
recorded from Madison (Montgomery, 1932) and Orleans parishes

150 Tulane Studies in Zoology Vol. 1

(von Reizenstein, 1863; Jung, 1939, 1950).

Phoebis sennae sennae (Linne). — This subspecies is included
on the basis of Klots' (1951) mention that "in southern Florida and
Louisiana many very sennae-like specimens occur." He describes the
color as being a warmer yellow than that of eubule. Indeed, in the
Tulane and Southwestern Louisiana Institute collections there are
specimens of P. sennae that fit this description, particularly those from
the Lafayette area. Since the subspecific classification is very com-
plex and for the most part a matter of statistics the writer has chosen
to include the "sennae-like" specimens in the records for eubule. One
female taken at Lafayette on November 2, 1950 comes closest to
fitting the description of warm yellow.

Phoebis philea (Linne), the Red-barred Sulphur. — The writer
has never collected P. philea although several specimens from Louisi-
ana have been available for examination. In addition to these the
writer has information from Dr. Wm. T. M. Forbes (in personal
communication) that he had taken this species in New Orleans on
January 1, 1933. The species is known to be migratory.

P. philea has not been recorded from Louisiana previously. The
few records are: Allen: Kinder, VI- 12 -41, (SLI); Orleans: New Or-
leans, X-ll-32, 1 $ (TU P-978), and VII-15-32, 1 5 (TU P-982).

Phoebis agarithe maxima (Neumoegen), the Large Orange
Sulphur. — As this butterfly is mainly tropical, only three specimens
have been caught in Louisiana, and these are from the southern
parishes. The species is migratory according to Williams (1937).

It has not been recorded from the state previously. The few records
are: Lafayette: Lafayette, 1 5 (SLI); Orleans: New Orleans, VIII-
24-26, 1 2 (TU P-965), IX-19-31, 1 2 (TU P-977), and X-2-31,.
1 2 (TU P-978).

Eurema daira DAIRA (Latreille), the Fairy Yellow. — This is a
very variable species; some have all, others only one-half the border
of the hind wings black, and individuals of the latter type often have
the undersides of the hind wings white. These two forms for a long
time were held as two species, but Dr. R. L. Chermock (personal
correspondence) considers them to be variations of one species.

In Louisiana the species apparently is not widespread, nor does it
fly during as great a part of the year as does E. lisa. With the ex-
ception of one male from the southwest, all specimens were taken
during the summer in the Florida parishes. Though the species ap-
parently is not evenly distributed, the writer found a dense population
of what appeared to be a southwesterly migration September 6-8,
1950, near Covington, La. All the butterflies were flying in one
direction a few feet from the ground stopping only occasionally to
feed at conspicuous flowers.

E. daira daira has not been recorded previously from Louisiana.

The records are: Lafayette: Lafayette, IX-25-36, 1 $ (SLI); Liv-
ingston: Springfield, IX-6-50, 2 $S, 1 2 (TU 2183, 2184); SL

No. 10 Lambremont: Butterflies and Skippers 151

Helena: Liverpool, IX-7-50, 3 $$,l 9 (TU 2193); Greensburg,
IX-7-50, 2 $ $ (TU 2194); St. Tammany: Folsom, IX-7-50, 1 $,
2 5 9 (TU 2187); Covington, IX-8-50, 2 $ $ (TU 2196); Tangi-
pahoa: Lee Landing, IX-6-50, 1 $ (TU 2181); Robert, IX-6-50,
1 $, 1 9 (TU 2185); Tangipahoa, IX-7-50, 1 9 (TU 2191);
Amite, IX-7-50, 2 9 9 (TU 2195); Washington: Angie, VI-5-50,
15,19 (TU 2034); Richardson, IX-7-50, 1 S,2 99 (TU 2188).

Eurema nicippe nicippe ( Cramer ). — The Nicippe Sulphur is
common throughout Louisiana during most of the summer months.
However, it has a relatively long flying season; earliest and latest
dates are April 8, 1950 (1 9 ) and December 8, 1949 (1 9 ).

This butterfly is not usually common everywhere throughout the
summer, but occurs in small swarms in sunny places in the pinelands.
When it is flying it is rare to see only one at a time, as usually
several are in flight short distances from each other.

Forty-eight specimens (30 $$, 18 99) have been examined
from 26 localities in the following parishes: Avoyelles (SLI), Cata-
houla, Claiborne, Concordia, Evangeline, Lafayette (SLI), Livingston,
Morehouse, Ouachita, Plaquemines, Pointe Coupee, Rapides, Richland,
Sabine, St. Helena, St. Landry (SLI), St. Tammany, Tangipahoa,
Vermilion (SLI) and Washington. Previously it had been recorded
from Madison (Montgomery, 1932) and Orleans parishes (von
Reizenstein, 1863).

Eurema Lisa (Boisduval and LeConte). — The Little Sulphur
is the most abundant pierid in the state. In Louisiana it is most
abundant in mid-summer, although it flies from early spring well
into the autumn. Earliest and latest dates are January 2, 1950
(2 9 9) and October 16, 1949 (4 S S,4 99). Scudder (1889)
stated that the butterfly will fly in heavily clouded weather, but the
writer believes this to be unlikely as it has always been observed to
take cover when clouds hide the sunlight.

E. lisa is not a strong flier and is easy to capture. Usually they fly
close to the ground in open places, and congregations of them are
often seen sucking water from moist ground. However, in spite of
its frailness and weak flight, it has been known to migrate in great
clouds for remarkable distances (Scudder, 1889). The larvae feed
on Cassia spp.

One hundred and eleven specimens (73 $ $ , 38 99) have been
examined from 56 localities in the following parishes: Allen, Assump-
tion, Bienville, Catahoula, Claiborne, Concordia, DeSoto, Evangeline,
Franklin, Lafayette (SLI), Lafourche, Livingston, Morehouse, Natchi-
toches, Orleans, Pointe Coupee, Rapides, Richland, Sabine, St. Bernard,
St. Charles, St. Helena, St. Landry, St. Tammany, Tangipahoa, Wash-
ington, Webster and West Feliciana. Previously it had been recorded
from Cameron (Hine, 1906), Madison (Montgomery, 1932) and
Orleans parishes (Jung, 1950).

Nathalis iole (Boisduval), the Dwarf Yellow. — Jung (1950)
reported this species as very common about fields in New Orleans,

152 Tulane Studies in Zoology Vol. 1

but I have not found it in great numbers at any one time as it is
very local and easily overlooked. It flies in sunny, grassy places along
the sides of roads, and in open fields. It is most abundant in the
warmer months, but stragglers have been taken as late as October in
Louisiana.

The few records are: Claiborne: 8 miles east of Lisbon, VIII-6-49,
1 $, 1 2 (TU 1457); Lafayette: Lafayette, X-2-36, 1 5 (SLI);
Pointe Coupee: McCrea, X- 14-49, 1 3 (TU 1568); Sabine: Negreet,
VII-7-50, 1 2 (TU 2101); 10 miles west of Negreet, VII-8-50,
1 2 (TU 2106).

Pieris RAPAE (Linne). — The Cabbage Butterfly was not origin-
ally a member of the North American fauna, but was introduced on
this continent at Quebec in I860 from Europe. From there the
species has spread throughout the continent inflicting damage to
crops totalling hundreds of thousands of dollars annually. Scudder
(1889) gave an excellent account of its spread, but gave no definite
date for its introduction into Louisiana. However, it had reached
Bastrop, Texas by 1881.

In Louisiana the species is abundant during the spring and early
summer months, but may be seen flying during late fall and winter.
Earliest and latest dates are January 2, 1950 (5 $$, I 2) and
November 30, 1950 (1 $). The writer has seen many adults feeding
on the blossoms of clover (Trifolium repens).

Twenty-one specimens (13 $ $ , 8 2 2) have been examined from
nine localities in the following parishes: Catahoula, Lafayette (SLI),
Ouachita, St. Charles, Washington and West Feliciana. Previously
it had been recorded from Orleans and St. Bernard parishes by Jung
(1939, 1950).

Pieris protodice protodice (Boisduval and LeConte). — The
Common White has a wide range over the continent and it is not
scarce in Louisiana where it flies during the hot summer months.
Earliest and latest dates are June 16, 1950 (5 $ $ , 1 2) and Sep-
tember 5, 193 8(1 2). The flight is easy and rather methodical, and
adults are easily collected.

Twelve specimens (8 $ $ , 4 22) have been examined from seven
localities in the following parishes: Catahoula, Concordia, Lafayette
(SLI), Morehouse, Orleans and Ouachita. Previously it had been
recorded from Cameron (Hine, 1906), Madison (Montgomery, 1932)
and Orleans parishes (von Reizenstein, 1863; Jung, 1939, 1950).

Ascia monuste phileta (Fabricius), the Great Southern
White. — Much has been written concerning the migration and im-
mature stages of this butterfly by Nielsen and Nielsen (1950) who
found eggs deposited on saltwort (Batis maritima), nasturtium
(Tropaeolum majus), spider-wisp (Cleome spp.) and other plants.
In spite of its wide range and abundance in the Gulf states only two
males and one female have been caught in the state. However, the
writer saw many of them flying along the shore and over the water
of Lake Pontchartrain at New Orleans (May 10, 1950), but speci-

No. 10 Lambremont: Butterflies and Skippers 153

mens were not obtained.

The few records are: Lafayette: Lafayette, VI-25-28, 1 5 (SLI)
and VII-8-50, 1 S (SLI); Orleans: New Orleans, VI-30-31, 1 &
(TU P-974). Previously it had been recorded from Orleans parish
by von Reizenstein (1863) and Jung (1939, 1950).

Superfamily HESPEROIDEA, the Skippers
Family Hesperiidae

Epargyreus clarus (Cramer), the Silverspotted Skipper. — The
flight of this species is very fast, and although it is common it is not
easily caught. All specimens were captured while they fed on flowers,
particularly blossoms of the buttonball bush (Cephalanthus occiden-
tal). Caterpillars feed on Robina, Wisteria, Pueraria and other
leguminous plants.

Jung (1950) reported it common within residential New Orleans,
but the writer has taken specimens only in sunny fields and bordering
areas of woods.

The few Louisiana records are: Caldwell: Columbia, VIII-3-48,
1 $ (TU 429); Sabine: Negreet, VII-7-50, 1 $ (TU 2101); St.
Tammany: Folsom, IX-7-50, 1 $ (TU 2187); Covington, IX-8-50,
1 $,l $ (TU2196). Previously it had been recorded from Madison
(Glick, 1939) and Orleans parishes (von Reizenstein, 1863; Jung,
1950).

Urbanus PROTEUS (Linne), the Longtailed Skipper. — In Louisiana
this species is seen almost throughout the year flying from flower to
flower in open sunny places. Earliest and latest dates are February
23, 1949 (1 S ) and November 6, 1949 (15). This skipper often
picks a resting place and returns to the same spot time and time
again after being disturbed. One such individual, whose tails were
broken in such a way as to make it easily recognizable, favored a lead
pipe in the writer's garden. Day after day for almost a month the
same skipper headquartered around the pipe, occasionally flying a
few feet away to feed on lantana flowers. The larvae feed on certain
species of the Leguminosae, particularly Wisteria and Clitoria.

Twelve specimens (10 2 2,2 52) have been examined from
eight localities in the following parishes: Allen, Assumption, Orleans,
Pointe Coupee, St. Bernard, St. Charles, St. Tammany and Tangi-
pahoa. Previously it had been recorded from Orleans parish by
Shufeldt (1884) and Jung (1950).

ACHALARUS LYCIADES (Geyer), the Hoary-edge. — Only three in-
dividuals have been taken in Louisiana, and these during the summer.
One was captured while it was feeding on flowers of the buttonball
bush (Cephalanthus occidentalis) . The larvae feed on various le-
guminous plants.

The species has not been recorded from Louisiana previously. The
few records are: Sabine: Negreet, VII-7-50, 1 $ (TU 2102); 10
miles west of Negreet, VII-8-50, 1 $ (TU 2106); Union: Marion,
VI-24-50, 1 $ (TU 2075).

154 Tttlane Studies in Zoology Vol. 1

Thorybes bathyllus (J. E. Smith). — The Southern Dusky-wing
is more abundant in Louisiana that its ally, T. confusis, and flies from
early spring throughout the summer. Earliest and latest dates are
February 24, 1950 (1 $) and September 8, 1950 (2 SS, 1 5).
The larvae feed on various species of the family Fabaceae.

T. bathyllus has not been recorded from Louisiana previously. The
ten specimens examined (6 $ $, 4 9 9) are from the following
localities: Bienville: Bienville (TU 1459); Sabine: Negreet (TU
2101); St. Tammany: Hickory (TU 1800); Covington (TU 2196);
Tangipahoa: Ponchatoula (TU 2182); Robert (TU 2185); Union:
7 miles east of Farmerville (TU 1863); Washington: Richardson
(TU 2188).

Thorybes confusis Bell, — The Northern Dusky-wing is found
occasionally in Louisiana during the summer months but records of
it are not numerous. According to Scudder (1889) it frequents open
fields and sunny places visiting flowers, often clovers. Little is known
of the immature stages.

T. confusis has not been recorded from Louisiana previously. The
few records are: Catahoula: 5 miles southwest of Lee Bayou, VI-11-50,
1 5, (TU 2044); Claiborne: Scottsville, VI-29-50, 1 S (TU 2086);
Sabine: Belmont, VIII-2-49, 1 $ (TU 1379); Negreet, VII-10-50,
1 $ (TU 2103).

Pyrgus communis communis (Grote). — The Common Checker-
spot Skipper is one of the commonest skippers of Louisiana, often
seen flying a few inches from the ground on hot, dusty roads. Earliest
and latest dates are January 24, 1950 (1 $ ) and October 16, 1949
(3 $ $). Most specimens have been taken during flight, which is
very rapid and elusive. The collector must not lose sight of the
insect for a second if he wishes to catch it, for its checkerboard pattern
blends very well with the background.

There is no apparent localization in the state, the species having
been collected in all the major areas. The larvae feed on various
species of the family Malvaceae.

Twenty-one specimens (18 $$, 3 2?) have been examined
from 18 localities in the following parishes: Catahoula, East Feliciana,
Evangeline, Jefferson, Lafayette (SLI), Natchitoches, Orleans, Oua-
chita, Pointe Coupee, Rapides, Sabine, St. Bernard, St. Charles, St.
Landry, St. Tammany, Washington and West Feliciana. Previously
it had been recorded from Orleans parish by Jung (1950).

Celotes nessus (Edwards), the Streaky Skipper. — The writer
has not seen this species in Louisiana but it was reported from
Madison parish by Glick (1939) who did extensive work with traps
carried by airplanes. A single specimen was taken during the day
at an altitude of twenty feet.

Pholisora hayhurstii (Edwards), Hayhurst's Skipper. — A sin-
gle specimen of this species has been taken in Louisiana, flying near
the ground in a dense oak-hardwood stand on the front lands of the

No. 10 Lambremont: Butterflies and Skippers 155

Mississippi river. Little is known concerning the immature stages.

The species has not been recorded from Louisiana previously. The
single record is: Jefferson: Kenner, III-9-51, 1 2 (TU 2298).

Erynnis horatius (Scudder and Burgess), Horace's Dusky-
wing. — Most of the individuals were captured in hot, sunny spots in
and along dense forests, but none were seen feeding. The flight is
rapid and fleeting, the insect staying about ten feet away from the
collector, alighting on the ground, and darting to a new resting place
if disturbed. Three males collected on June 4th, and 5th were sitting
on the hot, dry surface of a sand bar along the Pearl river. Earliest
and latest dates are June 1, 1950 ( 1 $ ) and September 7, 1950 (15).
Little is known about the immature stages.

Eleven specimens (9 $ $ ,2 2 2 ) were examined from nine local-
ities in the following parishes: Assumption, DeSoto, East Feliciana,
LaSalle, St. Tammany, Union and Washington. Previously it had
been recorded from Orleans parish by Jung (1950).

Erynnis juvenalis (Fabricius). — Only three specimens of this
skipper have been taken in Louisiana. The larvae feed on oaks and
various legumes according to Scudder (1889).

This species has not been recorded from Louisiana previously. The
records are: Orleans: New Orleans, II-4-50, 1 2 (TU 1782); St.
Tammany: Covington, IX-8-50, 2 2 2 (TU 2196).

Ancyloxypha numitor (Fabricius). — The Numitor Skipper-
ling apparently is never abundant in Louisiana. According to Scudder
(1889) the flight is slow and feeble and it is one of the easiest
skippers to capture in flight. The larvae feed on grasses (Klots,
1951). Earliest and latest dates are February 18, 1950 (1 S) and
October 19, 1949 ( 1 $ ) .

Eight specimens (3 $ $ , 5 $9) have been examined from seven
localities in the following parishes: Allen, Evangeline, Jefferson, St.
Bernard, St. Charles, St. Tammany and Washington. Previously it
had been recorded from Jefferson parish by Jung (1950).

Copaeodes minima ( Edwards ), the Tiny Skipperling. — This is
probably the smallest skipper in North America. It is very common
but its size and ability to blend into its surroundings make it difficult
to see. The majority were taken while feeding on daisies in open,
sunny fields throughout the state. Earliest and latest dates are March
2, 1949 and October 18, 1949- Nothing is known of the early
stages.

C. minima has not been recorded from Louisiana previously.
Twenty-four specimens (13 $$, 11 $2) have been examined
from 14 localities in the following parishes: Claiborne, Evangeline,
Jefferson, Livingston, Morehouse, Orleans, Ouachita, Pointe Coupee,
Rapides, Richland, St. Bernard and Tangipahoa.

Hesperia leonardus stallingsi H. A. Freeman. — A single
specimen of the Leonardus Skipper was taken in Madison parish by
Glick (1939) in an airplane trap at an altitude of twenty feet. The

156 Tulane Studies in Zoology Vol. 1

writer has no other information concerning the occurrence of this
species in Louisiana.

Hylephila PHYLEUS (Drury). — The Fiery Skipper is often
taken while it frequents flowers along the borders of woods, and in
open places in dense forests. The flight is not rapid, hence the
species is easily captured. Earliest and latest dates are March 2,
1949 (1 5) and October 16, 1949 (1 $, 1 5). The larvae sup-
posedly feed on crab grass (Scudder, 1889).

Twenty-one specimens (13 $ $ , 8 5 5) have been examined from
15 localities in the following parishes: Catahoula, East Feliciana,
Evangeline, Jefferson, Morehouse, Orleans, Richland, Sabine, St.
Landry, Tangipahoa, Union and Washington. Previously it had been
recorded from Cameron parish by Hine (1904).

Atalopedes campestris (Boisduval), the Sachem. — The few
Louisiana specimens are all from the eastern parishes near the Mis-
sissippi river. From what the writer has observed the flight is rather
rapid and jerky, the skipper flitting from flower to flower; however,
when it is feeding it may be caught easily. The larvae feed on
Bermuda grass.

The species has not been recorded from Louisiana previously. The
records are: East Feliciana: Felixville, VI-8-50, 2 9 5 (TU 2039);
LaSalle: Summerville, VI-12-50, 1 $ (TU 2048); Morehouse: 10
miles west of Twin Oaks, VI-26-50, 14,15 (TU 2079); Ouachita:
Perryville, VI-23-50, 1 $ (TU 2071); Sterlington, VI-18-50, 1 $
(TU 2060) and VI-23-50, 1 5 (TU 2072); Union: Marion, VI-
24-50, 15,15 (TU 2075).

POLITES MANATAAQUA MANATAAQUA (SCUDDERJ, the Manata-
aqua Skipper. — Lindsey, Bell and Williams (1931) stated that this
subspecies occurs from the northeastern United States to Alabama,
and westward to South Dakota and Nebraska. Klots (1951) described
its range as above, but included Arkansas and Georgia. The species
is rare in Louisiana, but nevertheless the six specimens taken con-
stitute what appears to be a southern range extension. All were
captured in moist, grassy openings in the pine flats of the Florida
parishes north of Lake Pontchartrain. The larvae feed on common
grasses.

The subspecies has not been recorded from Louisiana previously.
The records are: St. Tammany: Hickory, IX-13-50, 1 $ (TU 2198);
Tangipahoa: Ponchatoula, IX-6-50, 4 $ $,l 5 (TU 2182).

Polites THEMISTOCLES (Latreille). — The Tawny-edged Skipper
is well distributed throughout Louisiana. All specimens thus far
have been collected during the summer months in open sunny spots
in fields and woods, especially where there are flowers in bloom. The
caterpillars feed on grass.

The species has not been recorded from Louisiana previously. The
records are: East Feliciana: Felixville, VI-8-50, 1 $ (TU 2039);
Sabine: Negreet, VII-7-50, 1 5 (TU 2101); St, Tammany: Hickory,

No. 10 Lambremont: Butterflies and Skippers 157

VII-15-50, 1 9 (TU 2001); Tangipahoa: Lee Landing, IX-6-50,
1 $, 1 9 (TU 2181); Ponchatoula, IX-6-50, 1 $ (TU 2182);
Union: Haile, VI-20-50, 1 $ (TU 2067).

POLITES vibex BRETTUS (Boisduval AND LeConte), the Whirl-
about.— There is no apparent localization of this species in Louisiana,
and although it is not common, it has been taken often in the pine
flats of the Florida parishes and in other parts of the state. All speci-
mens were collected while they were feeding on wild flowers. The
larvae feed on grasses.

P. vibex brettus has not been recorded from Louisiana previously.
The records are: Evangeline: 12 miles west of Mamou, X- 16-49, 1 9
(TU 1576); Sabine: Negreet, VI-7-50, 2 4 $, 2 9 9 (TU 2101);
8 miles west of Negreet, VII-8-50, 1 $ (TU 2107); 5"/. Tammany:
Covington, IX-8-50, 1 $ (TU 2180) and IX-8-50, 3 $ $ (TU
2196); Tangipahoa: Robert, IX-6-50, 2 $$ (TU 2185); Union:
Haile, VI-20-50, 1 9 (TU 2067).

Wallengrenia otho otho (J. E. Smith), Otho's Skipper. —
Thus far in Louisiana this species has been collected mostly in the
Florida parishes. The larvae feed on crab grass (Panicum sanguinale)
according to Klots (1951).

Otho's skipper has not been recorded from Louisiana previously.
The records are: Orleans: New Orleans, 111-22-50, 1 $ (TU 1821);
St. Tammany: Covington, 4 $ $, 2 9 9 (TU2196); Tangipahoa:
Ponchatoula, IX-6-50, 1 $ (TU 2182).

Poanes viator ( Edwards ), the Broadwinged Skipper. — Although
this species apparently is locally very common in Louisiana during
the spring and early summer months, only twelve specimens have
been collected thus far. All of these were taken together while they
were feeding on a roadside strip of daisies. The flight is exceptionally
fast. Lindsey, Bell and Williams (1931) stated that the species flies
from June to August. Nothing is known of the early stages.

The Louisiana record is: St. Charles: Norco, 111-10-51, 6 $ $,
6 9 9 (TU 2301). Previously it had been recorded from Orleans
parish by Scudder (1889).

Atrytone arogos arogos (Boisduval and LeConte), the
Iowa Skipper. — A single specimen has been collected in Louisiana
while it was feeding on daisies growing in a garbage dump. Little
information is available on the immature stages.

The species has not been recorded from Louisiana previously. The
lone record is: St. Tammany: Covington, IX-8-50, 1 $ (TU 2197).

Atrytone dukesi Lindsey, Dukes' Skipper. — A single specimen
has been taken in Louisiana. The species was described from
Mobile, Alabama, and Holland (1931) predicted that it would occur
over a wider area along the Gulf, although Klots (1951) made no
mention of it west of Alabama. The Louisiana record constitutes a
possible westward range extension. Information is lacking concern-
ing the early stages.

158 Tulane Studies in Zoology Vol. 1

The species has not been recorded from Louisiana previously. The
lone record is: West Carroll: 11 Vi miles west of Oak Grove, VI-19-50,
1 $ (TU2064).

Atrytone ruricola metacomet (Harris). — The Dun Skipper
apparently is not common in Louisiana, as only two specimens have
been taken, and those were in the western and northern part of the
state. Almost nothing is known about the immature stages of this
skipper.

As ruricola metacomet has not been recorded from Louisiana pre-
viously. The records are: Sabine: Negreet, VII-7-50, 1 S (TU
2101); Union: Marion, VI-24-50, 1 $ (TU 2075).

Oligoria maculata (Edwards), the Twin Spot. — Edwards
(1865) described this species from New Orleans, but both Holland
(1931) and Lindsey, Bell and Williams (1931) considered it to be
most abundant toward the Carolinas. Klots (1951) stated that it
strays west to Texas. Indeed, the few specimens collected in Louisi-
ana have all been taken from the eastern part of the state, particularly
in the pine flats of the Florida parishes. Very little is known con-
cerning the immature stages, although the larvae presumably feed on
grass (Klots, 1951).

The records are: St. Tammany: St. Tammany, IV-13-50, 1 9 (TU
1903); Covington, IX-8-50, 1 $ (TU 2196); Hickory, IX-13-50,
1 $ (TU 2198); Tangipahoa: Ponchatoula, IX-6-50, 2 $ $ (TU
2182).

Lerema accius (J. E. Smith). — The Accius Skipper apparently
is not abundant in Louisiana. Thus far it has been taken during the
late summer months only, although it has been reported to fly during
all months of the year in the southernmost limits of its range (Lindsey,
Bell and Williams, 1931). The larvae feed on grasses and Indian
corn according to Klots (1951).

The records are: Pointe Coupee: McCrea, X- 14-49, 1 4,1 ? (TU
1567, 1568); St. Tammany: Covington, IX-5-50, 1 5 (TU 2180)
and IX-8-50, 3 $ $, 1 2 (TU 2196); Folsom, IX-7-50, 1 $ (TU
2187); Tangipahoa: Ponchatoula, IX-6-50, 1 $ (TU 2182). Pre-
viously it had been recorded from Madison (Glick, 1939) and Orleans
parishes (Edwards, 1865).

Amblyscirtes vialis (Edwards). — A single specimen of the
Roadside Skipper has been taken in Louisiana. Klots (1951) stated
that the range is from southern Canada to Florida, and westward to
Texas, but Holland (1931) wrote that it was common in the Mis-
sissippi valley. Lindsey, Bell and Williams (1931) stated that vialis
apparently is replaced by another species in the southeastern states.
There can be no doubt, however, that vialis is an occasional visitor in
Louisiana, particularly in the northern parishes. The caterpillars feed
on grasses.

A. vialis has not been recorded from Louisiana previously. The
record is: Webster: Leton, VIII-6-49, 1 $ (TU 1449).

No. 10 Lambremont: Butterflies and Skippers 159

Amblyscirtes textor (Hubner). — The Wovenwinged Skipper
is represented in Louisiana by a single specimen. This individual was
taken while it was feeding on a blackberry flower. Nothing is known
of the immature stages.

A. textor has not been recorded from Louisiana previously. The
record is: Jefferson: Kenner, III-9-51, 1 $ (TU 2297).

Lerodea l'herminieri (Latreille), the Fuscous Skipper. — Five
specimens of this uncommon skipper have been taken in Louisiana
during the summer months. Little is known about the immature
stages.

L. l'herminieri has not been recorded from Louisiana previously.
The records are: St. Helena: Liverpool, IX-7-50, 1 $ (TU 2193);
Tangipahoa: Ponchatoula, IX-6-50, 3 $ $ (TU 2182); Union: Haile,
VI-20-50, 1 $ (TU 2066).

Lerodea eufala (Edwards), the Eufala Skipper. — This species
also is rare in Louisiana. Only four specimens have been taken in the
state and these in late summer and early autumn. The larvae feed
on grasses.

The records are: St. Bernard: Shell Beach, X- 18-49, 2 $ $,19 (TU
1595); Tangipahoa: Lee Landing, IX-6-50, 1 9 (TU 2182). Pre-
viously it had been recorded from Madison parish by Glick (1939).

CALPODES ETHLIUS (Stoll), the Brazilian Skipper. — Although
this species apparently is common in Louisiana, only one specimen
has been taken during the survey. The flight is very rapid, the skipper
rising and falling in a very quick undulating manner.

Jung (1950) reported large numbers of the larvae on canna in
New Orleans, and the single adult specimen in the Tulane collection
was reared from a pupa found on canna leaves. Dr. Forbes informed
the writer (in correspondence) that he observed numerous larvae and
adults of ethlius in New Orleans on January 1, 1933.

The single record is: Orleans: New Orleans, X- 17-50, 1 2 (TU
2224).

Panoquina panoquin (Scudder). — The Salt-marsh Skipper has
not been taken by the writer in Louisiana, but is included here on
the basis of Scudder's (1889) report of it from New Orleans. Noth-
ing is known about the early stages.

Panoquina ocola (Edwards). — The Ocola Skipper is fairly
common in Louisiana during the late summer, frequenting the edges
of forested areas. Nothing is known about the immature stages.

P. ocola has not been recorded from Louisiana previously. The
records are: Allen: Oberlin, X-16-49, 1 ? (TU 1574); Hampton,
X-16-49, 1 $ (TU 1579); Evangeline: St. Landry, X-14-49, 1 9
(TU 1569); Point Blue, X-16-49, 1 $ (TU 1579); Pointe Coupee:
McCrea, X-14-49, 15,1$ (TU 1567, 1568); St. Bernard: Delacroix
Island, X-18-49, 1 $ (TU 1592); St. Charles: Norco, 1-24-50, 1 S
(TU 1763); St. Tammany: Folsom, IX-7-50, 1 5 (TU 2187);
Tangipahoa: Ponchatoula, IX-6-50, 2 $ $,\ $ (TU 2182).

160 Tulane Studies in Zoology Vol. 1

Summary
Butterflies and skippers were collected in Louisiana during three
years (1948-1951) and these records are coupled with those of other
institutional and published records to form a composite list for the
state. Before the writer's survey was begun there had been 43 species
recorded, and during its progress a local study by Jung (1950) added
17 new state records, bringing the total to 60 by July 1950. With
the completion of the present study 38 previously unrecorded species
are added, and the state list now totals 98.

An attempt was made to collect in many areas of the state, and
during different times of the year. Fifty of the 64 parishes of
Louisiana were sampled, and two additional ones are included in the
literature. Best represented are the southeastern parishes, New Orleans
and vicinity. A total of 1501 adult butterflies and skippers were
examined in the course of the study.

Each species on record is listed with observations by the writer and
a summary of the collection data arranged by parishes.

From laboratory analyses of the records certain geographic patterns
are noted. While most species are found distributed more-or-less
evenly throughout the state, some are restricted to particular physio-
graphic or vegetational regions. Euptychia areolata areolata, E. gemma
gemma, Celastrina argiolus pseudargiolus, Eurema daira daira, Wall-
engrenia otho otho and Oligoria maculata appear to inhabit only the
Florida Parish pine area, with E. gemma gemma occasionally visiting
the Upland pine areas. Euptychia cymela cymela, Anaea andria,
Graphium marcellus marcellus and Anthocharis genutia are more-or-
less restricted to upland pine and hardwood areas. Heliconius char-
it onius, Agraulis vanillae nigrior, Euptoieta claudia claudia and
Papilio cresphontes cresphontes are restricted almost entirely to the
southern parishes.

An analysis of the flying dates leads to the general conclusion that
most of the common species fly almost throughout the year in Louisi-
ana, with their peaks of abundance in the hot summer months. Figure
3 summarizes the seasonal distribution of all of the commoner species.

On comparison with published distribution patterns it appears that
the Louisiana specimens of Melitaea nycteis nycteis constitute a range
extension of significance. Since the species is not abundant in Louisi-
ana this must be the southern tip of its range. The collections of
Polites manataaqua manataaqua, Hemiargus isolus and Atrytone dukesi
assist in defining the ranges of these species.

The 98 species recorded here are not all of the butterflies and
skippers expected from the state. More concentrated sampling over
a longer period of the year, and a more intense consideration of the
immature stages will undoubtedly reveal a greater number of species.
Perhaps in due time the writer may be able to fill in the vacancies,
however, it is hoped that this paper will stimulate entomologists to
initiate further studies of the Lepidoptera of Louisiana.

No. 10

Lambremont: Butterflies and Skippers

161

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Figure 3. Summary of flying seasons of the commoner species in
Louisiana; thickened parts of bars indicate months of greatest
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Acknowledgements
The writer wishes to express his thanks and sincere appreciation
to Dr. George H. Penn, Tulane University, without whose encourage-
ment this research could not have been completed. Thanks are due
to Dr. E. S. Hathaway, Tulane, for the interest he showed during
this study, and to Dr. Ralph L. Chermock, University of Alabama,

162 Tulane Studies in Zoology Vol. 1

who so willingly identified the specimens sent to him for confirmation,
and who criticized the preliminary and final drafts of the manuscript.
Appreciation is due to Dr. George H. Bick, Tulane, for helpful sug-
gestions and for collections from the Delta region. The writer is
indebted to Dr. Fred R. Cagle, Tulane, for invitations extended to
accompany him in the field. Dr. Lewis T. Graham kindly placed the
collections of Southwestern Louisiana Institute at the writer's disposal.
Invaluable aid was rendered by Dr. W. T. M. Forbes, Cornell Uni-
versity, and by Dr. Austin H. Clark, U. S. National Museum, both of
whom carefully criticized the manuscript. Also, to his wife who spent
many hours preparing the final draft the writer will be ever grateful.

References Cited

Beall, Geoffrey 1946. Seasonal variation in sex proportions and

wing length in the migrant butterfly, Danaus plexippus L. 3.

Migrants of the autumn in Louisiana. Trans. Royal Ent. Soc.

London, 97: 342-343.
1948. The fat content of a butterfly, Danaus

plexippus Linn., as affected by migration. Ecology, 27: 80-94.
Chermock, Ralph L. 1947. A new subspecies of Limenitis archip-

pus. Amer. Mus. Novitates, No. 1365: 1-2.
Clark, Austin H. 1941. Notes on some North and Middle American

danaid butterflies. Proc. U. S. Nat. Mus., 90: 531-542.
[Comstock, John H.] 1882. The hog caterpillar of the orange. Rept.

U. S. Comm. Agric., 1881: 246-248.

Comstock, William P. and F. Martin Brown 1950. Geographical
variation and subspeciation in Heliconius charitonius Linnaeus.
Amer. Mus. Novitates, No. 1467: 1-21.

Dos Passos, Cyril F. 1938. A new race of Basilarchia archippus
Cramer from Louisiana. Canad. Ent., 70: 243.

Edwards, William H. 1865. Description of certain species of di-
urnal Lepidoptera found within the limits of the United States
and British America. Proc. Ent. Soc. Phila., 4: 201-204.

Field, William D. 1941. Additional notes on Calycopis cecrops
(Fabricius) and Calycopis beon (Cramer). Jour. Kans. Ent.
Soc, 14: 66-69.

Floyd, Ernest H. 1941. Investigations on the biology and control
of the alfalfa caterpillar, Colias eurytheme iBoisduval. La. Agric.
Exp. Sta., Louisiana Bull., No. 323: 16-25.

Gerould, John H. 1946. Hybridization and female albinism in
Colias philodice and Colias eurytheme. A New Hampshire sur-
vey in 1943 with subsequent data. Ann. Ent. Soc. Amer., 39:
383-397.

Glick, P. A. 1939. The distribution of insects, spiders, and mites in
the air. U. S. Dept. Agric, Tech. Bull., No. 673 : 1-150.

Hine, James A. 1904. A contribution to the entomology of the re-
gion of the Gulf Biologic Station. Gulf Biol. Sta. Bull., No. 2:
65-68.

1906. A second contribution to the entomology of

the region of the Gulf Biologic Station. Gulf Biol. Sta. Bull,
No. 6: 65-83.

Holland, William J. 1931. The Butterfly Book, revised edition.
New York: Doubleday and Co., pp. 1-424.

No. 10 Lambremont: Butterflies and Skippers 163

Hovanitz, William 1943. The nomenclature of the Colias chryso-

theme complex in North America. Amer. Mus. Novitates, No.

1240: 1-4.
1944. The ecological significance of the color

phases of Colias chrysotheme in North America. Ecology, 25:

45-60.

Howard, L. O. 1895. The gray hair-streak butterfly and its damage
to beans (Uranotes melinus Hubn.). Insect Life, 7:354-355.

Johnson, Frank and William P. Comstock 1941. Anaea of the
Antilles and their continental relationships, with descriptions of
new species, subspecies and forms. Jour. N. Y. Ent. Soc, 49 :
301-343.

Johnson, Lawrence 1886. [Communication]. Proc. Ent. Soc. Wash.,
1: 34-35.

Judd, W. W. 1950. An assemblage of monarch butterflies (Danaus
p. plexippus) on the north shore of Lake Erie. Jour. N. Y. Ent.
Soc, 58: 169-172.

Jung, Rodney C. 1939. On the occurrence of some of southern
Louisiana's more common butterflies. Tulane Biol., 3 : 17-18.
(mimeographed) .

1950. An annotated list of the Lepidoptera of the

New Orleans area. Proc. La. Acad. Sci., 8: 42-48.

Klots, Alexander B. 1951. A Field Guide to the Butterflies of
North America, East of the Great Plains. Boston: Houghton
Mifflin Co., pp. 1-349.

Kopman, Henry H. 1903. The butterflies of Louisiana. New Or-
leans Times Democrat, April 12, 1903.

Lindsey, A. W., E. L. Bell and R. C. Williams 1931. The Hesper-
oidea of North America. Denison Univ. Bull., Jour. Sci. Lab-
oratories, 26: 1-142.

McDunnough, J. 1938. Checklist of the Lepidoptera of Canada
and the United States of America, Parts I and II. Mem. So.
Calif. Acad. Sci., 1: 1-272.

Morris, John G. 1862. Synopsis of the described Lepidoptera of
North America. Smithso7i. Misc. Collections, 4: 1-358.

Michener, C. D. 1947. A review of the subspecies of Agraulis
vanillae L. Amer. Mus. Novitates, No. 1215: 1-7.

Montgomery, Robert W. 1932. Records of Louisiana butterflies.
Ent. News, 43: 182.

Nielsen, Erik T. and Astrid T. Nielsen 1950. Contributions to-
ward the knowledge of the migrations of butterflies. Amer.
Mus. Novitates, No. 1471: 1-29.

Park, Orlando 1948. Observations on the migrations of monarch
butterflies through Evanston, Illinois in September, 1948. Chi-
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[Riley, C. V. and L. O. Howard] 1895. A new food plant for Pap-
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Rothschild, Walter and Karl Jordan 1906. A revision of the Am-
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Saunders, William 1871. On the swarming of Danaus archippus.
Canad. Ent., 3:156-157.

Saverner, P. A. 1908. Migrating butterflies. Ent. News, 19:
218-220.

164 Tulane Studies in Zoology Vol. 1

Scudder, Samuel H. 1889. The Butterflies of the Eastern United
States and Canada with Special Reference to New England.
Cambridge: published by the author; vols. I-III.

Shufeldt, R. W. 1884. Observations upon a collection of insects
in the vicinity of New Orleans, La., during the years 1882 and
1883. Proc. U. S. Nat. Mus., 7: 331-338.

Skinner, Henry 1897. A new species of Debis. Ent. News, 8 : 236.

1907. Studies of Thecla irus Godart and Thecla

henrici Grote and Robinson. Ent. News, 18: 129-132.

Viosca, Percy Jr. 1933. Louisiana Out-of-Doors. New Orleans:
published by the author, pp. 1-187.

von Reizenstein, Ludwig 1863. Catalogue of the Lepidoptera of
New Orleans and its Vicinity. New Orleans: Isaac T. Hinton,
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568-585.

— /V r\ - /V / *-w v^t/^=»c*4«i

sR- 3 1961

hkebsity

Volume 1, Number 11

August 26, 1954

TWO NEW SPECIES OF THE GENUS GRAPTEMYS

FRED R. CAGLE,

DEPARTMENT OF ZOOLOGY, TULANE UNIVERSITY,
NEW ORLEANS.

TULANE UNIVERSITY
NEW ORLEANS

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Hobart M. Smith, Professor of Zoology, University of Illinois,
Urbana, Illinois.

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of Florida, Gainesville, Florida.

T. Paul Maslin, Associate Professor of Biology, University of
Colorado, Boulder, Colorado.

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written copy, double-spaced, and carefully corrected.

Separate numbers may be purchased by individuals, but subscriptions
are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
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When citing this series authors are requested to use the following
abbreviations: Tulane Stud. Zool.

Price for this number: $0.75.

Assistants to the Editor:
Carol L. Freret
Robert E. Gordon

George Henry Penn, Editor,
c/o Department of Zoology,
Tulane University,
New Orleans, U. S. A.

HARVARD
UNIVERSITY

TWO NEW SPECIES OF THE GENUS GRAPTEMYS
FRED R. CAGLE,

Department of Zoology, Tulane University,
New Orleans.

Two distinctive, undescribed populations of turtles were found in
Mississippi and Alabama by the Tulane University field crews of 1952
and 1953. The turtles are abundant at the type localities and are the
dominant species of turtles present.

Both species are closely related to Graptemys oculifera (Baur) and
with this species form a unique complex of three allopatric species
ranging from the Pearl River in the west to the Alabama river in the
east. The three groups of populations are recognized as species
because: (a) the degree of difference between the three groups is
much greater than that between the subspecies of related species oc-
curring in the central United States; (b) the degree of difference is
as great as that occurring between sympatric species of this and
related genera; (c) there are no intermediates. These factors suggest
that these populations have been reproductively isolated for a long
period. It is not possible to conclude that these three forms would
not be capable of interbreeding if they were not spatially isolated.

The following abbreviations are used in the descriptions: CL, cara-
pace length (not along curve); Cw., carapace width; PL, plastron
length; Pw., plastron width; Hw., head width; Ht., height; Aw.,
alveolar width. All measurements are maximum. The size of in-
dividuals is expressed as plastron length.

UMMZ = University of Michigan Museum of Zoology; CNHM r=
Chicago Natural History Museum.

The illustrations in color were provided through the cooperation
of Roger and Isabelle Hunt Conant who had photographed the turtles
for inclusion upon one of the color plates in their forthcoming Field
Guide to the Reptiles and Amphibians. Mr. George P. Meade gener-
ously provided the funds for their publication here. The research
was aided by a grant from the National Science Foundation.

The collections essential to these descriptions could not have been
made without the enthusiastic field work of graduate students. I am
especially grateful to A. H. Chaney, Robert Gordon, Donald Tinkle,
Robert Webb, Walter Stone, and Howard Suzuki.

GRAPTEMYS FLAVIMACULATA, sp. nov.

Yellow-blotched Sawback

Holotype. — Tulane 14798, a juvenile male, collected from the Pas-
cagoula River, 13 miles S.W. of Lucedale, George Co., Mississippi,
18 Aug. 1952 (figs. 3, 5, 7).

Paratypes.—Tuhne 14752, 14754, 14756-766, 14768, 14772, 14774-
776, 14778-785, 14788, 14790, 14795, 14799-802, 14804, 14806-809,
14811-812, 14815, 14818, 14821-822, 14825-827, 14829, 14832-833,
14842, 14845-846, 14850, 14852-854, 14857-858, 14862-863, 14865-

168 Tulane Studies in Zoology Vol. 1

871, 14873-875, 14920-921, 14935, 14938; CNHM 69806-808;
UMMZ 108567-571, 36 juveniles, 44 males and 3 females collected
at the type locality by a Tulane University field crew.

Hypodigm, — A total of 124 preserved specimens, 74 examined by
dissection and not retained, and many others observed in the field
provided a series of observations for the analysis of variation of this
species. The study of these individuals has also provided a basis for
determination of some aspects of the behavior and for estimating the
population structure.

Diagnosis. — Graptemys jlavimaculata is a member of a narrow-head
complex that includes three species occurring only in Louisiana, Mis-
sissippi and Alabama, Graptemys oculifera in the Pearl River drain-
age, Graptemys jlavimaculata in the Pascagoula River and its tribu-
taries, Graptemys nigrinoda in the Black Warrior and Alabama Rivers

(fig. 9). '

G. jlavimaculata differs from G. nigrinoda in that: the postorbital
mark is rectangular or triangular, is broader than any of the lateral
neck lines entering the orbit, and is usually joined to a longitudinal
line; the ventral surface of the jaw is dominated by broad yellow
lines much wider than the olive-green interspaces; each costal with
a large yellow or orange central blotch or crescent; the vertebral spines
are high, narrow, laterally compressed; the edge of the carapace of
males and juveniles is much less serrate (Table 1).

G. jlavimaculata differs from G. oculijera in that: the orbital mark
is broad and is ordinarily joined to a dorsal longitudinal neck line;
the lower jaw is dominated by broad yellow lines; each costal has a
large blotch or crescent of yellow.

Description of holotype — Carapace length, 8.05 cm; carapace width
at juncture of fourth and fifth marginals, 6.15 cm; carapace width at
juncture of seventh and eighth marginals, 6.72 cm; plastron length,
7.43 cm; width of posterior lobe of plastron, 3.63 cm; height (meas-
ured in vertical line through spine of second vertebral), 3.70 cm; height
at spine of third vertebral, 3-41 cm; head width (measured at anterior
edge of tympanum) 1.24 cm; length of symphysis of lower jaw, 0.44
cm; alveolar width of upper jaw, 0.25 cm.

Edge of carapace serrate; each marginal projecting beyond anterior
corner of the next posterior one. Each marginal with a wide, yellow
bar or semicircle of yellow. Each costal with a broad ring or yellow
blotch (figs. 1,3).

Plane of the plastron 1.40 cm below a plane through the edge of
the fifth to the seventh marginals. Yellow color predominant on the
plastron, bridge and ventral surface of marginals. Black plastral marks
extending along the sutures between each pair of shields ( fig. 7 ) .

Neck with 19 longitudinal, yellow lines; those on the ventral sur-
face twice the width of those on the dorsal surface. Longitudinal,
interorbital line nearly as wide as widest dorsal neck lines and termi-
nating above postorbital mark. Three yellow lines between the orbits.
Two broad, yellow lines entering the orbit below a triangular post-

No. 11

Cagle: Two New Graptemys

169

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170 Tulane Studies in Zoology Vol. 1

orbital mark (figs. 1, 5).

Horny edges of both upper and lower jaws light in color. Ventral
surface of lower jaw with two broad, transverse, yellow bands.

Anterior surface of forelegs with wide, yellow lines; those from
the base of second and fourth toes are the broadest.

Description of paratypes. — The paratypic series is divided into juve-
niles, adult males and adult females to insure adequate description.

Juveniles. — The juveniles represent individuals in the first to the
tenth seasons of growth (figs. 11, 13). The color pattern of the
individuals greater than 9 cm in plastron length is basically the same
as that of the smaller juveniles but the contrast between the yellow
or orange markings and the background is less.

The head pattern in lateral view consists of two wide, yellow lines
extending posteriorly from the orbit along the neck, a triangular or
vertically elongated postorbital mark, and a broad yellow line extend-
ing the length of the lower jaw. The postorbital mark is, in most
individuals, joined to a longitudinal line on the dorsal surface of the
head (figs. 1, 11, 13).

The head pattern from the dorsal view consists of a wide yellow
line between the orbits extending from a point just above the nostrils
to above the center of the postorbital mark; two lines from the upper
edge of the orbit that, in most individuals, join the postorbital mark
and continue posteriorly onto the neck. The latter lines are inter-
rupted in some individuals. The background color is black (figs.
11,13,15).

The lower jaw is dominated by wide yellow lines broadly bordered
with black. Of the three or four transverse yellow lines, the most
posterior ones join longitudinal lines (figs. 11, 13, 15). The neck
has 17-20 wide yellow lines around its circumference at the rear of
the skull.

The carapace has bright yellow or orange central splotches on each
costal. These are extremely variable in shape but usually cover more
than one-third the total area of each shield. In some individuals
(14795) they are rounded, in others (14788) they are crescent-like.
The upper surface of each marginal has a semicircle of yellow en-
closing a darker area having a faint indication of two concentric light
lines. Each of the vertebral spines is conspicuously colored black.

The plastron has a pattern of black lines tending to extend along
the seams. The background is cream or light.

The anterior surface of the forelimbs has fine yellow lines on a
black background. Those extending from the base of the second and
fourth toes are the wider.

The jaws have a thin, sharp-edged horny covering. That of the
upper overlaps the lower to provide a scissor-like action. The
alveolar surfaces are not broadened in the larger turtles, Hw/Aw =
4.7-5.8; no correlation with size.

The head is narrower in relation to plastron length in the larger
juveniles. Pl/Hw = 4.1-4.5 in those 5.0 cm to 5.3 cm in plastron

No. 11 Cagle: Two New Graptemys 171

length; 5.5-6.6 in the larger turtles.

The carapace width decreases in proportion to plastron length in
the larger individuals; Pl/Cw = 0.91-0.99 in those 5.0 cm to 5.5 cm
in plastron length, 0.95-1.1 in the larger ones. The height is reduced
in larger turtles; Pl/Ht = 1.7-1.9 in those 5.0 cm to 5.5 cm in plastron
length; 1.8-2.1 in the larger ones.

The spines of the first four vertebrals are conspicuous in all the
juveniles (fig. 11). In those with a plastron length of less than 11
cm the height of the spines of the second vertebral is 2/3 to 3/4 the
length of the seam between the second and third marginal. The ele-
vations on the first and fourth vertebrals are ridge-like, those of the
second and third are distinct spines.

Adult males. — The adult* males in the paratypic series range from
7.10 cm to 9.5 cm in plastron length. The smaller adult males (less
than 8 cm in plastron length) have the same color pattern as the
juveniles. The larger males (more than 8 cm in plastron length)
lose the orange tint in the blotches of the carapace and the lines of
the head. The plastral pattern is lightened and poorly defined in
the larger individuals.

The spines of the vertebral shields are slightly lower in the large
males than in the juveniles and those of the first and third shields
are rounded (14850, 14858). The carapace edge of the largest male
is less serrate anteriorly than in smaller males; the posterior lateral
corners of the fifth and sixth marginals project but slightly beyond
the anterior lateral corner of the next posterior marginal (fig. 15).

The height is not reduced with increased length as it is in other
species, Pl/Ht = 1.9-2.2. The carapace is somewhat narrowed as the
male increases in length; Pl/Cw = 1.0-1.1.

The head is slightly narrower in relation to plastron length in
larger males, Pl/Hw = 5.6-5.8 in those 7-8 cm in plastron length;
Pl/Hw = 6.0-6.3 in those greater than 8 cm in length. There is no
consistent broadening of the alveolar surfaces in larger males, Hw/Aw
= 5.1-6.1.

The third claw of the forefoot is slightly longer than the length
of the seam between the second and third marginals.

Adult females. — The adult females retain the color pattern of the
juveniles but the blotches on the carapace and the black lines of the
plastron are not as distinct. The largest female (14815) has the
black background of the carapace lightened by the development of
irregular areas of olive-green. The black borders of the yellow marks
on the marginals are indistinct. The only remaining vestige of the
black lines of the plastron are present on the gular, humeral and anal
shields.

The spines of the vertebral shields are much more reduced than
in adult males but remain clearly evident on the first four vertebrals
(height of spine of second vertebral = 1/4-1/3 length of seam be-
tween first and second marginals). The carapace edge is not serrate
except between the eighth to the twelfth marginals.

172 Tulane Studies in Zoology Vol. 1

The smallest mature female (14920) in the paratypic series has
the following measurements: PI. 14.0, CI. 14.9, Cw. 13-0, Ht. 6.5,
Hw. 2.1, Aw. 0.43- The largest (14815) has these measurements:
PI. 15.9, CI. 17.4, Cw. 14.3, Ht. 7.3, Hw. 2.33, Aw. 0.52.

The carapace height is not reduced in comparison with the juve-
niles and adult males, Pl/Ht = 1.9-2.2. The carapace is slightly-
broadened in contrast with the adult males, Pl/Cw = 1.1 (in all three
individuals).

The head is proportionately broader than in adult males, Pl/Hw =
4.5-4.9 and the alveolar surfaces are wider, Hw/Aw = 4.5-4.9. The
third claw of the forefoot is 1/2 to 2/3 the length of the seam between
the second and the third marginal.

Range. — Specimens are available from three localities; the type
locality, the Chickasawhay River and 15 miles from the mouth of the
Pascagoula River, Jackson County, Mississippi (AMNH 46774, col-
lected by M. J. Allen, 1930). All specimens have been taken from
the Pascagoula River or its tributaries. It is apparent that this species
is restricted to this stream system as repeated efforts to collect it
from adjacent rivers to the east and west were unsuccessful (fig. 9).
Populations undoubtedly occur in all the major tributaries and should
be especially abundant in the Escatawpa River.

Habitat. — The Pascagoula River basin, comprising 8,900 square
miles, has had the greatest concentration of industry in Mississippi.
Two of the tributaries, Tallahala and Sowashee Creeks, have been
highly polluted by sewage and waste from wood pulp and chemical
industries (Anderson, 1950). This pollution has apparently had no
tendency to reduce the Graptemys populations and has probably,
through contributing to the increase of the mollusc populations, im-
proved the general habitat for G\ jlavimaculata.

The river at the type locality has a sand and clay bottom and many
sand bars and beaches. Flood-stranded debris is abundant and much
of the shoreline has a stand of dense brush. The debris and brush
extending into the water provides shelter for turtles. The river is
75 to 250 feet in width and deep pools are separated by long stretches
of shallow water. A moderately rapid current maintains a narrow
channel through the shallow areas.

These turtles were most abundant about the tangled roots of trees.
Often several turtles were observed lying together in such places.
The approach of the boat usually caused them to leave the resting
site and swim rapidly toward deep water.

The Chickasawhay River is 30-150 feet wide; the banks have
exposed areas of limestone in many places and the bottom is generally
rock and sand. Deep, quiet pools are connected by fast-flowing
stretches of shallow water. Drift wood and rocks are abundant along
the shores. No G. jlavimaculata were taken in the faster current or
the static water but they were abundant in the snags in the slower
current.

This species occurs with Graptemys pulchra, Pseudemys floridana

No. 11 Cagle: Two New Graptemys 173

mobilensis, Pseudemys scripta elegans and Sternotherus carinatus. It
is clearly the dominant turtle species in the Pascagoula and Chicka-
sawhay Rivers.

Bionomics. — G. flavimaculata is a comparatively small, rapidly
growing turtle adapted to live in moderate current. The smallest
sexually mature male studied had a plastron length of 6.67 cm; the
smallest mature female had a plastron length of 13-30 cm.

The males may reach sexual maturity in the second growing season.
The smallest mature male was in its second growing season when col-
lected and three others ranging in size from 7.47 cm to 8.00 cm were in
the third and fourth seasons. The largest male studied had a plastron
length of 10.95 cm.

The age at maturity of females could not be determined as the
early growth rings were not evident in any mature individual. None
of the females larger than 14 cm in length had measurable growth
zones.

GRAPTEMYS NIGRINODA, sp. nov.

Black-knobbed Sawback

Holotype. — Tulane 14662, a juvenile female, collected from the
Black Warrior River, above Lock 9, 17.5 miles SSW of Tuscaloosa,
Tuscaloosa County, Alabama (figs. 4, 6, 8).

Paratypes.— Tulane 14643, 14647-648, 14652-653, 14655-657,
14659, 14664-665, 14682, 14691, 14694, 14697, 14700, 14706,
14708-710, 14714, 14720, 14723, 14725, 14729; CNHM 69809-811,
UMMZ 108572-574, 26 juveniles, 5 adult males.

Diagnosis. — Graptemys nigrinoda differs from G. flavimaculata
and G. oculifera in that: the yellow postorbital mark is boomerang-
shaped and is joined to the mark of the opposite side by a diagonal
line; the interorbital line is narrow or indistinct; each costal has a
circular or semicircular mark formed by a narrow yellow line; the
vertebral spines are flattened, compressed, knob-like; the carapace is
very serrate.

Hypodigm. — A total of 117 individuals were available for study
from two localities and numerous others were observed in the field.
The data derived from field observations and a study of the 117
preserved individuals provide the basis for the information presented.

Description of the holotype. — Maximum carapace length, 8.02 cm;
carapace width at juncture of fourth and fifth marginals 6.54 cm;
carapace width at juncture of seventh and eighth marginals 6.91 cm;
maximum plastron length 7.52 cm; maximum width of posterior lobe
of plastron, 3.87 cm; maximum height (measured in vertical line
through spine of second vertebral, 3.42 cm; height at spine of third
vertebral 3.30 cm; maximum head width (measured at anterior edge
of tympanum) 1.32 cm; length of symphysis of lower jaw 0.57 cm;
alveolar width of upper jaw 0.28 cm.

Edge of carapace very serrate (fig. 4). General color of carapace
dark olive green. Faintly defined, narrow circular or semicircular
mark formed by a thin yellow line on each marginal and costal (figs.

174 Tulane Studies in Zoology Vol. 1

2,4).

Plane of the plastron 1.23 cm below a plane through the edge of
the fifth to the seventh marginals. Yellow color predominant on
plastron and ventral surfaces of marginals. Branching, black pattern
on plastron (fig. 8).

Neck with 25 longitudinal yellow lines; those on the ventral surface
not much wider than those on the dorsal surface. Longitudinal inter-
orbital line narrower than lateral neck lines. Fine yellow lines be-
tween orbits. Four yellow lines entering the orbit laterally; two of
these broader than others. Postorbital mark boomerang-shaped and
joined to mark of opposite side by a diagonal line ( fig. 2 ) .

Ventral surface of lower jaw with a broad, transverse yellow line
(fig. 8).

Anterior surface of forelegs with five yellow lines.

Description of the paratypes. — The paratypic series is divided into
two groups, juveniles and adult males.

Juveniles. — The juveniles are in their first or second season of
growth. The background color of the carapace is dark olive green
and that of the soft parts intense black.

The head pattern differs from that of juveniles of flavimaculata
in that the postorbital yellow mark is a vertically elongated, boomerang-
shaped line extending posteriorad on the dorsal surface to join the
mark of the opposite side to form a Y. The interorbital, longitudinal
stripe is narrow and indistinct in many individuals.

The lower jaw has one broad, yellow transverse line sometimes
bordered both anteriorly and posteriorly by narrow yellow lines. Two
of the wide, longitudinal neck lines join behind the level of the jaw
juncture to form another transverse line. The lower surface of the
neck is dominated by black on which the yellow lines appear to be
superimposed (figs. 10, 12).

The carapace has a complete circle of a narrow yellow line on each
costal. The knob-like vertebral spines are intense black. The upper
surface of each marginal has a semicircular mark formed by a narrow
yellow line. The ventral surface of each marginal may be yellow
with an ocellus of black and yellow rings, or the yellow may be re-
duced to an irregular blotch on the medial side with the remainder
of the marginal covered with alternating black and yellow lines
(figs. 2, 8).

The plastron is yellow to bright orange with black lines extending
along the seams (fig. 8). The anterior surface of the forelimb has
two to five yellow lines. When four or five are present, those from
the base of the second and fourth digits are much the wider.

The jaws are as described for flavimaculata. The alveolar width is
contained 4.0-6.0 times in the head width. There is no relative
broadening of the alveolar surface in the larger individuals.

The head is narrower in relation to plastron length in the larger
turtles than in the smaller, Pl/Hw = 4.6-5.6. The carapace width
decreases in proportion to plastron length with size, Pl/Cw = 0.9-1.0.

No. 11

Cagle: Two New Graptemys

175

u
o

S
to

to .

fcJ3

176

Tulane Studies in Zoology

Vol. 1

Figure 3. Dorsal view, G. flavimaculata, sp. nov. (Photograph by
Isabelle Hunt Conant).

Figure 4. Dorsal view, G. nigrinoda, sp. nov. (Photograph by Isabelle
Hunt Conant).

No. 11

Cagle: Tivo New Graptemys

111

Figure 5. Lateral view, G. flavimaculata, sp. nov. (Photograph by
Isabelle Hunt Conant).

■
MR

Figure 6. Lateral view, G. nigrinoda, sp. nov. (Photograph by Isa-
belle Hunt Conant).

178

Tulane Studies in Zoology

Vol. 1

Figure 7. Ventrai view, G. flavimaculata, sp .nov. (Photograph by
Isabelle Hunt Conant).

jBh,.^

14662 «_^>

Figure 8. Ventral view, G. nigrinoda, sp. nov. (Photograph by Isa-
belle Hunt Conant).

No. 11 Cagle: Tivo New Graptemys 179

The height is proportionately reduced in the larger turtles Pl/Ht =
1.8-2.2.

The spines of the first four vertebrals are high (height of second
equals 2/3 or more lengths of seam below second and third marginal)
and are much broadened and rounded. This is in sharp contrast with
jlavimaculata.

Adult males. — The adult males range from 6.80 cm to 8.00 cm in
length. They have the same color pattern as the juveniles but the
yellow markings of the carapace and the black markings of the plastron
are more obscure. The lines of the head in the largest male ( 14648 )
are fainter and the black background is lightened.

The spines of the vertebral shields are worn and eroded in the
largest specimen and do not have the broadened, rounded appearance
of the spines of juveniles. The carapace edge is less serrate than in
the four smaller individuals (fig. 14).

The height is reduced in the largest male in relation to plastron
length (Pl/Ht = 2.0-2.2 in smaller animals; 2.3 in largest). This
is primarily a reflection of the worn spines. The carapace is not
narrower in the larger males, Pl/Cw = 0.9-1.0; in four smaller ones,
Pl/Cw =1.0 in largest.

The head is slightly narrower in relation to plastron length in the
larger males; Pl/Hw = 5.3-5.5 in those 6.0 cm-7.1 cm in plastron
length, Pl/Hw = 5.5.-6.0 in those greater than 7.5 cm in plastron
length. There is a slight broadening of the alveolar surfaces in rela-
tion to head width, Hw/Aw = 5.8 in smallest, 4.8 in largest.

The third claw of the forefoot is slightly shorter than the length of
the seam between the second and third marginal.

Adult females. — No adult females were collected but one was
studied with binoculars and a 20 power telescope in the field. This
large female retained the distinctive markings of the juveniles. The
black background color of the head and the brilliant yellow markings
were not reduced.

Range. — Specimens were collected from only two localities, the type
locality and the Alabama River 5.5 miles east of Gosport, Monroe
County, Alabama. Collecting on the Coosa River and the Mulberry
Fork of the Black Warrior River did not yield this species. Its absence
from these streams poses the problem of what limiting factors have
prevented its movement northward. An attempt to approximate the
northernmost occurrence in the Black Warrior was made June 18.
A count of basking turtles was made from the juncture of Locust
Fork and Mulberry Fork to the Barney Coal Mine during the period
from dawn until 1 PM. No G. nigrinoda were observed until the
river widened approximately three miles above the Barney mine.
Basking turtles were common along the entire stretch. There was an
apparent correlation between the occurrence of G. nigrinoda and the
occurrence of Amyda. Graptemys pulcbra is present in Mulberry
Fork and Locust Fork.

Habitat. — Generally similar to that of G. oculijera and G. jlavima-
culata. All individuals were taken from brush or logs in current or

180

Tulane Studies in Zoology

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90 85

Figure 9. The distribution of G. oculifera (narrow vertical lines),
G. flavimaculata (horizontal lines), and G. nigrinoda (wide ver-
tical lines). The black spots indicate collecting sites; the circles
indicate localities where intensive collecting has failed to pro-
duce members of this complex.

were shot from basking sites above current. Both the Black Warrior
and Alabama Rivers are wider, deeper rivers than the Pascagoula or
the Pearl Rivers but they also have sand and clay bottoms and moderate
current in those areas where G. nigrinoda was collected.

This species occurs in association with G. pulchra, Pseudemys flori-
dana mobilensis, Pseudemys scripta elegans, Sternotherus carinatus, and
Amyda sp.

Bionomics. — The smallest male has a plastron length of 6.80 cm
and is in the third season of growth. The largest male observed was
8.76 cm in length and had no growth zones present. No adult
females were collected.

DISCUSSION

Other turtle species occurring in the Pearl, Pascagoula and the lower
parts of the Tombigbee and Alabama rivers exhibit a gradual coast-
wise gradient in most of their characters. The populations of Pseu-
demys scripta, Pseudemys floridana, Kinosternon subrubrum, Amyda
ferox and Graptemys pulchra of each river system are different in
the percent expression of characters but much overlap is evident.

Graptemys pulchra occurs in these same stream systems as well as
in the Escambia to the east and the upper reaches of the Alabama
where G. nigrinoda is absent. However, the populations of the upper
Alabama are conspicuously different from those of the lower. This

No. 11 Cagle: Two New Graptemys 181

difference in parts of the Alabama is also reflected in the fact that
Graptemys geographica occurs in the upper reaches but not in the
lower.

The restriction of G. nigrinoda to that part of the river below the
fall line may be suggestive of a rather recent, rapid evolution of this
complex since the exposure of the Gulf Coast in Pliocene time. There
is a correlation between the northern limit of distribution and the
isocryme of 15° C (as reported by Hedgpeth, 1953).

The absence of these three forms in river systems east of the Ala-
bama and west of the Pearl is further evidence that the evolutionary
history of this complex is substantially different from that of other
turtle species of the Gulf Coast (fig. 9). Unfortunately there is but
limited information available on the history of these river systems.
The extent of the Alabama and Tombigbee systems as contrasted
with the Pearl and Pascagoula is suggestive of greater age. Perhaps
in this situation rests the explanation of the greater degree of diverg-
ence in G. nigrinoda, it perhaps being the older of the three forms.
The biologist has no precise knowledge of comparative rates of evolu-
tion in the turtles but their long fossil record suggests unusually slow
response. The relative youth of the Gulf Coast streams implies that
rapid evolution has occurred in this complex.

These current-dwelling turtles do not inhabit the extreme head
waters and smaller tributaries as do members of the genera Amyda,
Pseudemys and Kinosternon. There are no reports of overland move-
ments although such observations are common for other turtles. Their
isolation is thus possibly much more complete than is implied by the
proximity of the drainage systems. The cohesiveness of the three
groups would suggest the designation of a superspecies. This would
serve to emphasize the differences between these and the remainder
of the genus. Each of the three groups of populations could be desig-
nated a semispecies (Mayr, 1942; Cain, 1953).

Although the three forms are basically similar in their osteology
and general configuration, the morphological gap between G. nigri-
noda and G. jlavimaculata is much greater than that between G.
jlavimaculata and G. oculifera (Cagle, 1953b). Particularly strik-
ing is the contrast between the flattened carapace and knob-like
projections in G. nigrinoda and the elevated carapace and laterally
compressed spines of the other two species. An alternative taxonomic
arrangement would be the recognition of G. jlavimaculata as a sub-
species of G. oculifera while retaining G. nigrinoda as a separate
entity. Such an arrangement would place emphasis on the differences
in degree of divergence between the forms but the evolutionary
implications of such a grouping can not be justified.

Related species of this genus do not have such striking differenti-
ation in adjacent stream systems. Graptemys geographica, Graptemys
pseudogeographica and Graptemys kohni occupy more extensive
ranges without developing such gaps or steepening gradients in vari-
ation (Cagle, 1953a).

182

Tulane Studies in Zoology

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No. 11 Cagle: Two New Graptemys 183

A TENTATIVE KEY TO THE GENUS GRAPTEMYS
This key is presented as an aid to identification of the members of
this genus. It is not intended to present those characters that are of
fundamental significance in establishing- the patterns of speciation.

1. Length of symphysis of lower jaw %

longer than minimum distance be-
tween the orbits and a small triangu-
lar, yellow spot back of orbit sepa-
rated from orbit by 2 to 3 diagonal
yellow lines Graptemys geographica (LeSueur)

(Geographic turtle. Mississippi and
St. Lawrence basins. Northern Lou-
isiana north through eastern Oklaho-
ma and Kansas to northern Minne-
sota; eastward through Missouri,
Illinois, Kentucky, Tennessee to Ver-
mont. Northern Mississippi, Ala-
bama.)

Length of symphysis of lower jaw equal
to or less than minimum distance be-
tween the orbits; or, if symphysis is
longer than minimum distance be-
tween orbits, no triangular spot be-
hind eye but a large, irregular-
shaped greenish or yellowish post-
orbital blotch present 2

2. Each costal with a large yellow-orange

blotch, crescentic mark or ring. 3

Costals without markings as described 5

3. Each costal with a large yellow or

orange blotch or crescentic mark cov-
ering much of the shield. Yellow the
dominant color on the ventral surface
of jaw and neck Graptemys flavimaculata n. sp.

(Yellow -blotched sawback. Pasca-
goula River and tributaries, Missis-
sippi.)

No such markings 4

4. Each costal with one complete circle of

yellow or orange formed by a line
broader than the widest neck line. A
yellow postorbital ovoid, rectangular
or rounded spot not connected with a
dorsal, longitudinal line. Vertebral
spines laterally compressed Graptemys oculifera (Baur)

Figures 10-15. Juveniles and males of G. nigrinoda and G. flavimacu-
lata; scale line = 5 cm. 10, Hatchling, G. nigrinoda; 11, Indi-
vidual in first season of growth, G. flavimaculata; 12, Juvenile
male, G. nigrinoda; 13, Juvenile male, G. flavimaculata; 14,
Adult male, G. nigrinoda; 15, Adult male, G. flavimaculata.

184 Tulane Studies in Zoology Vol- 1

(Ringed sawback. Pearl River and
tributaries, Louisiana and Missis-
sippi.)

Each costal with a yellow or orange
semicircle or circle formed by a line
much narrower than the widest neck
line. A yellow, vertical curved post-
orbital bar connected by a diagonal
line with the postorbital bar of the
opposite side. Vertebral spines knob-
like Graptemys nigrinoda n. sp.

( Black-knobbed sawback. Alabama
and Black Warrior River systems.)

5. Females never developing a head as
wide as Vs the carapace width. Post-
orbital mark not a large irregular
shaped greenish or yellow blotch —

Females developing a much broadened
head as wide as Vz the carapace
width. Postorbital mark an irregular
bordered greenish or yellowish blotch
or mark absent (see illustrations,
Cagle, 1952.) 10

Without vertebral spines (each vertebral
may be slightly elevated above next
posterior one). Maximum plastron
length about 16 cm. Graptemys versa Stejneger

(Texas map turtle. Colorado River
System, Texas.)

With distinct vertebral spines (absent in
some individuals greater than 16 cm
in plastron length.) — 7

7. A postorbital vertical line extending
from a dorsal' longitudinal line to the
base of the orbit and anteriorly
under the orbit; vertical line fre-
quently interrupted by a longitudinal
line or bar Graptemys kohni (Baur)

(Mississippi map turtle. Eastern
Texas, Oklahoma, Kansas and Ne-
braska east to Mississippi River and
southern Indiana.)

No postorbital vertical line; a comma-
shaped, rectangular or ovoid post-
orbital mark. A series of longitudi-
nal neck lines entering orbit below
postorbital mark. _ 8

No. 11 Cagle: Two New Graptemys 185

8. Postorbital spot, oval or elongate, much
smaller than orbit, and surrounded
by 2 or 3 concentric lines; 3 or 4 con-
spicuous longitudinal lines entering
orbit; these lines alternating with
thinner, less conspicuous lines. A
total of 6-7 lines entering orbit; a
wide yellow band extending the width
of the ventral surface of the jaw; no

large yellow spot beneath eye

Graptemys pseudogeographica sabinensis Cagle

(Sabine map turtle. Sabine River
of western Louisiana and eastern
Texas.)

Not as above. 9

9. Postorbital spot not elongate but rectan-

gular; length of rectangle extending
dorso-ventrally ; maximum of 5 longi-
tudinal lines entering the eye. No
wide yellow band extending the width
of ventral surface of jaw but a large
spot present at symphysis of lower
jaw; a large yellow spot beneath eye.
Graptemys pseudogeographica ouachitensis Cagle

(Ouachita map turtle. Louisiana
northwestward into eastern Oklaho-
ma, Kansas, Nebraska and eastward
through southern Wisconsin, Illinois
and southern Indiana to northern
Alabama. Eastward in the Ohio River
System.)

Postorbital spot comma-shaped; not rec-
tangular; maximum of 5 longitudinal
lines entering the eye ; no wide yellow
band extending the width of the ven-
tral surface of the jaw; no large spot
present on symphysis of lower jaw;
no large yellow spot beneath the eye.
Graptemys pseudogeographica pseudogeographica (Gray)

(False map turtle. Eastern Nebras-
ka, southeastern South Dakota east-
ward through southern Minnesota
and Iowa to eastern Illinois.)

10. An irregular bordered yellow bar ex-

tending completely across ventral
surface of lower jaw; margins of
plastral shields not bordered with
black Graptemys barbouri Carr

(Barbour's map turtle. Southwest-
ern Georgia and Panhandle of Flori-
da.)

186 Tulane Studies in Zoology Vol. 1

No bar as described ( an elongated blotch
or bar may be present) ; margins of
plastral shields bordered with black

(Alabama map turtle. Pearl River,
Louisiana eastward to Alabama,
Coosa and Escambia rivers.)

Graptemys pulchra Baur

REFERENCES CITED

Anderson, Irving E. 1950. Surface waters of Mississippi. Miss.
State Geol. Surv., Bull. 68: 1-338.

Cagle, Fred R. 1952. The status of the turtles Graptemys pulchra
Baur and Graptemys barbouri Carr and Marchand with notes on
their natural history. Copeia, 1952 (4) : 223-234.

1953a. Two new subspecies of Graptemys pseudo-

geographica. Occ. Pap. Mus. Zool. Univ. Mich., No. 546: 1-17.

. 1953b. The status of Graptemys oculifera. Zoolo-

gica, 38: 137-144.

Cain, A. J. 1953. Geography, ecology and coexistence in relation to
the biological definition of the species. Evolution, 7(1) : 76-83.

Hedgpeth, Joel W. 1953. An introduction to the zoogeography of
the northwestern Gulf of Mexico with reference to the inverte-
brate fauna. Publ. Inst. Marine Sci., 3(1) : 107-224.

Mayr, E. 1942. Systematics and the Origin of Species. New York,
Columbia Univ. Press, pp. 1-334.

O - /V/f -/V /'**" ~X^CL*^> I

MAR "3 1961

HARVARD
UNIVERSITY

■

Volume 1, Number 12

August 26, 1954

THE TAXONOMIC STATUS OF THE MID-GULF COAST

AMPHIUMA

IMOGENE R. HILL,

DEPARTMENT OF BIOLOGY, UNIVERSITY OF NEW MEXICO,
ALBUQUERQUE, NEW MEXICO.

TULANE UNIVERSITY
NEW ORLEANS

Manuscripts submitted for publication are evaluated by the editor
and by an editorial committee selected for each paper. Contributors
need not be members of the Tulane University faculty.

EDITORIAL COMMITTEE FOR THIS NUMBER

Arnold B. Grobman, Director, Florida State Museum, Gaines-
ville, Florida.

Laurence M. Klauber, Curator of Reptiles, Zoological Society
of San Diego, San Diego, California.

Robert C. Stebbins, Curator of Herpetology, Museum of Verte-
brate Zoology, University of California, Berkeley, Cali-
fornia.

Manuscripts should be submitted on good paper, as original type-
written copy, double-spaced, and carefully corrected.

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are not accepted. Authors may obtain copies for personal use at cost.

Address all communications concerning exchanges, manuscripts, edi-
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mittances should be made payable to Tulane University.

When citing this series authors are requested to use the following
abbreviations: Tulane Stud. Zool.

Price for this number: $0.35.

George Henry Penn, Editor,
c/o Department of Zoology,
Tulane University,
New Orleans, U. S. A.
Assistants to the Editor:

Carol L. Freret

Robert E. Gordon

MUS. C8MP. ZOOL
LIBRARY

MAR- 3 1961
Jast OL

THE TAXONOMIC STATUS OF THE MID-GULE

AMPHIUMA J

IMOGENE R. HILL,

Department of Biology, University of New Mexico,
Albuquerque, New Mexico.

The identity of the Amphiuma of the Gulf Coast has been studied
by Ryder (1880), Cope (1886), Brimley (1909), Goin (1938) and
Baker (1947). These investigators have drawn different conclusions
from the study of similar material. Their conclusions have been based
on analyses utilizing either one or several of the following characters:
number of toes, color, ratio of total length to body length, number of
costal grooves and ratio of body length to limb lengths.

The early investigators, Ryder, Cope and Brimley, had only a small
number of individuals available for their studies. The more recent
workers, Goin and Baker, considered populations of individuals, but
none of the investigators, measured or dealt with ontogenetic vari-
ation, allometric growth or sexual dimorphism. Since some authors
give only the mean of the ratios and of the costal groove number, it
is not possible to evaluate the characters reported. A new approach
was required to evaluate the characters previously used and to investi-
gate other differences, if any, between the populations.

Historical — Garden described Amphiuma means in 1821. In 1827
Cuvier described a three-toed individual as Amphiuma tridactylum
and compared it with a two-toed one: "This description of the
Amphiuma of three digits agrees nearly in everything to the kind
with two digits. Its general form is the same; its diameter is one-
twentieth of its length; the length of its head is nearly one-fourteenth;
its tail is exactly one-fourth. It has similar folds on each side. Its
color is the same. Summarizing, it is necessary for descriptive purposes
to look closely at its extremities. Then one sees that each of its limbs
is divided into three perfectly distinct toes; it is in reality its only
exterior character which is readily discernable."

The three-toed individuals were placed in the genus Mauraenopsis
by Fitzinger (1843). Ryder (1880) studied a series of young
Amphiuma from Biloxi, Mississippi and reestablished the three-toed
salamander in the genus Amphiuma. Ryder based his classification
solely on the number of digits. He stated "so that in some the
number of digits (two) is characteristic of Amphiuma, and in others
(three) they are characteristic of Mauraenopsis." Cope (1886) could
not separate his specimens into two-toed and three-toed groups and
so he classified all individuals, regardless of the number of toes, as
Amphiuma means.

Brimley (1909) called the three-toed animal Amphiuma tridacty-
lum.. Brimley stated that the three-toed specimens differed from the

1 From a thesis submitted in partial fulfillment of the require-
ments for the degree of Master of Science of the Graduate School
of Tulane University.

192 Tulane Studies in Zoology Y°^_*

two-toed ones in being lighter in color on both the dorsal and ventral
surfaces and in being distinctly bicolored; the limbs of the three-toed
Amphiuma were "more strongly developed." The character used in
separating the two forms, prior to 1909, was number of toes.
Brimley also reported that he had never seen a two-toed individual
which had attained as great a size as the three-toed ones; one three-
toed individual examined by Brimley was 736.6 mm in length.

Goin (1938) examined 206 individuals and found that the costal
groove count and the ratio of total length to body length may also
be used as taxonomic characters. Considering the region where the
two ranges meet, he stated "coloration, costal groove count and ratio
of total length to body length change more or less gradually, but
the two-toed specimens of the east interdigitate with the three-toed
specimens of the Mississippi Valley." According to Goin, Amphiuma
means means is a two-toed, unicolored animal with 57-60 costal
grooves and a ratio of total length to body length which averages
1.33. Amphiuma means tridactylum is a three-toed, bicolored animal
with 60-64 costal grooves and a ratio of total length to body length
which averages 1.29- Goin considered the area of intergradation as
the region between Lake Pontchartrain in southeastern Louisiana and
Liberty County, Florida.

Baker (1947) examined over 280 Amphiuma and concluded that
"two species of Amphiumae, Amphiuma means and Amphiuma tri-
dactylum, occur, each occupying a distinct ecological range." Baker
found no evidence of intergrades and stated that all known specimens
could be designated as one of these species by the "valid species
characters of number of toes, body length in total length, length of
limbs in body length and coloration."

Procedures

Number of toes and coloration — In the course of this investigation,
537 Amphiuma were examined. Two hundred and forty-six of these
were two-toed, two hundred and eighty-eight were three-toed, and
three individuals had both two- and three-toed limbs (Table 1). All
individuals examined are placed into one of six arbitrary geographical
series to determine the degree of difference, if any, between the
series. Series 1 includes all three-toed Amphiuma from the west
bank of the Mississippi River westward. The most western locality
of an Amphiuma examined in this study is nine miles northwest of
Saratoga, Hardin County, Texas. Series 2 consists of the individuals
from Audubon Park in New Orleans, Louisiana. Series 3 includes
all individuals in the alluvial valley east of the Mississippi River.
Included in Series 4 are the three-toed Amphiuma from the region
east of the alluvial valley of the Mississippi River to western Alabama.
This series includes individuals from extreme southeastern Louisiana
and Mississippi.

Series 5 consists of the two-toed individuals from the same geo-
graphical area as Series 4, and Series 6 consists of those individuals
from the Atlantic coastal states westward to Liberty County, Florida

No. 12

Hill: Taxonomic Status of Amphiuma

193

TABLE 1.
Number of Toes

Series

Number

with
Two Toes

Number

with
Uncertain
Number of

Toes

Number

with

Three Toes

Total

Number

Examined

191

1 9.9

163

Total

246

288

537

Figure 1. Distribution of the 537 Amphiuma examined in this re-
search. The circular and X-shaped symbols represent three-toed and
two-toed individuals respectively.

194 Tulane Studies in Zoology Vol. 1

(figure 1). The individuals from western Alabama to Liberty
County, Florida, were not placed into any of the six series because
they were too few in number to constitute an additional series. Series
4 and 5 extend as far east as the locality where both the two-toed and
the three-toed individuals are found in close association. This ar-
bitrary grouping into series does not mean to imply that the individ-
uals from western Alabama to Liberty County, Florida, are different
from those in Series 4 and 5.

Most of the specimens utilized in this study are deposited in the
collections of Tulane University (TU); a few were borrowed from
the American Museum of Natural History, the United States National
Museum, and the University of Florida.

An external morphological study was made on each individual.
The following information was recorded: museum catalogue number,
date collected, collector, tail length, body length, hind-limb length,
fore-limb length, head length, number of toes, sex and coloration.
The length of the tail was taken as the distance from the anterior
end of the cloacal opening to the tip of the tail. The limbs were
pressed posteriorly and horizontally along the body and the lengths
were recorded as the distances from the base of the limb to the
distal end of the longest digit. In the three-toed individuals, the
longest digit is the middle one and the outermost toe is the smallest
(when the limbs are pressed posteriorly and horizontally along the
body). The measurements were made consistently on the right limbs
of the animals unless the right limbs were mutilated. The head
length was taken as the straight line distance from the anterior end
of the brachial fissure to the tip of the nose. The number of toes
on each limb was recorded in a formula arrangement, e.g. 2/2 2/2.

The formula is read from left to right, and the numbers represent,
respectively, the number of toes on the right fore-limb, the number
of toes on the left fore-limb, the number of toes on the right hind-
limb and the number of toes on the left hind-limb. If the individuals
were sexed, this was done by dissection. The dorsal, ventral and
throat colorations were recorded. Table 2 indicates the throat colora-
tion of the specimens from the six geographical series.

The measurements were made with vernier calipers, except when
the structure was greater than 125 mm. In the latter case or if the
preserved animal were coiled, a string was used and transferred to a
ruler calibrated in millimeters. The measurements were recorded to
the nearest millimeter. All of the measurements were made on pre-
served specimens. There is undoubtedly some differential shrinkage
between the various body parts, resulting from the preservation, but
this possible error is likely to be minimized since all of the individuals
examined were preserved by comparable methods.

Only those individuals were measured whose parts were not muti-
lated. Mutilation could usually be detected by the presence of scar
tissue. Frequently, Amphiuma have part of a limb or an entire limb
missing. Obviously, it is easier to detect a mutilation which has

No. 12

Hill: Taxonomic Status

of Ampbiuma

195

TABLE 2.
Throat Coloration

Series

Number

without

Throat

Patch

Number

with

Uncertain

Throat Color

Number
with

Throat
Patch

Total

Number

Examined

191

1 3^

163

Total

246

288

537

occurred recently in the life of an animal than it is to determine one
which occurred very early in the life of the individual. It is unusual
to find all four limbs of an animal mutilated. The hind-limbs are
normally longer than the fore-limbs, except in individuals less than
a hundred millimeters in length. One can usually confirm the opinion
of whether or not a digital mutilation has occurred by comparing the
measurements of all four limbs.

Body proportions — Heretofore, three assumptions have been made
by some workers who have considered body proportions as taxonomic
characters in Ampbiuma. These assumptions include absence of
sexual dimorphism, high correlation between the dimensions of the
body parts, no ontogenetic variation. The latter is illustrated by
Baker's use of simple ratios, implying that body proportions do not
change with an increase in age (body length).

If these assumptions represent the facts, then the previous work is
valid, but if one or more of the assumptions are incorrect, the results
may require some qualification. Statistical methods can often be
used to give quantitative estimates of these assumptions. The pre-
liminary procedure was to determine the presence or absence of
sexual dimorphism, the degree of correlation between the body parts,
and whether or not ontogenetic variation was present.

To determine the presence or absence of sexual dimorphism, four
groups of graphs were set up, each group being composed of six
graphs representing the six geographical series. The four groups
represent the following four body proportions: head length to body
length, fore-limb length to body length, hind-limb length to body
length and tail length to body length. For each graph the horizontal
scale represents the body length in millimeters and the vertical scale
represents the other body part. The presence or absence of important
sexual dimorphism can be estimated by inspection. An inspection
of the scattergrams indicates that the males and females do not fall

196

Tulane Studies in Zoology

Vol. 1

o e

* x

cut©

©»

X 0

/•

Figure 2. Scattergrams showing the relationship of head length to
body length for each of the six series. The three symbols, X, circle
with dot and plain dot, identify respectively male, female and unsexed
individuals.

into two separate groups in any of the series; therefore, if sexual
dimorphism exists it is negligible ( figures 2-5 ) .

Obviously, there is a definite relationship of head length, fore-limb
length, hind-limb length and tail length to body length so the problem
is concerned with the degree of correlation and the nature of the
correlation (e.g., simple and linear). The degree of correlation of

No. 12

Hill: Taxonomic Status of Amphiuma

197

e>
©

S "
So

0«

•

•
•

M
x* «/»

X "

•

• •

0 «

•
•

•

x «

Figure 3. Scattergrams showing the relationship of fore-limb length
to body length for each of the six series. Symbol's as in fig. 2.

the various body parts with the body length was determined by cal-
culating the correlation coefficients (Tables 3-6). The method used
was Pearson's product-moment method using grouped data cross-
tabulated. Twenty-four correlation tables were constructed. In con-
structing the correlation tables, the body length was considered the
independent variable and the other body part length the dependent
variable (Peatman, 1947).

Ontogenetic variation — To determine whether or not body pro-

198

Tulane Studies in Zoology

Vol. 1

••

* X

9 X *
9

It 2

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Ok 0

X
9 O

* *

• •

X ^

* 9 *
X ©

X
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9
9
9

X
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X
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Figure 4. Scattergrams showing the relationship of hind-limb length
to bo dylength for each of the six series. Symbols as in fig. 2.

portions remain constant as the animal increases in age (length),
the regression lines for the four body parts of each of the six series
were calculated. The correlations in ontogenetic series are nearly
always high, but inspection of the scattergrams indicate that linearity
may be assumed for practical purposes.

The fact that the line is linear for the four body proportions of
each of the six series was established by the product-moment corre-
lation coefficients. Thus, the regression equation for the straight

line is:

X = a -\- aB

No. 12

Hill: Taxonomic Status of Ampbiuma

199

x°©
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* X

• •
©

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Figure 5. Scattergrams showing the relationship of tail length to
body length for each of the six series. Symbols as in fig. 2.

where X is equal to either head length, fore-limb length, hind-limb
length or tail length; a and b are the regression constants and B is
the body length corresponding to a particular X length. In order to
determine X for any particular body length, B, the regression con-
stants must be calculated.

The three methods suggested by Klauber (1943) for calculating
the regression constants are: (1) analytical methods, (2) method of
least squares and (3) the graphical method. Of these, the latter was

200

Tulane Studies in Zoology

Vol. 1

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204 Tulane Studies in Zoology Vol. 1

chosen, the reason being that the same graphs used to determine
sexual dimorphism could be used to determine the regression constants.

After obtaininng a and b for each of the four body proportions for
each of the six series, these constants were substituted in their
respective regression equations.

From a study of the regression lines, it was determined that the
body proportions do not remain constant throughout life. The reason
for the foregoing statement is that a is not equal to zero (the
regression line does not pass through the origin) (Klauber, 1943).
However, it should be mentioned that in some cases a is almost equal
to zero; although, additional collections may or may not reduce a to
zero. If the proportion remains constant throughout life, the relation-
ship of the head length to the body length should be the same in the
adults as it is in the juveniles. If a juvenile with a body length of
50 mm is substituted in the regression equation (H = 1.034 -\-
0.119B), the value of H is equal to 6.984 mm. If an adult body
length of 600 mm is substituted in the regression equation, H is
equal to 72.434 mm. The juvenile head length is 14 percent of its
body length; the adult head length is only 12 percent of its body
length. This indicates that the juveniles have proportionately larger
heads than the adults; therefore, the proportionality does not remain
constant with an increase in body length.

Standardization — Since there were insufficient numbers of individ-
uals.of a single body length in each series, a second method suggested
by Klauber was utilized to eliminate ontogenetic variation. This
method requires the conversion of the head length, fore-limb length,
hind-limb length, and tail length of the individual to a standard
body length. In using this method, the assumption was made that
any specimen in growing to (or returning to) the standard body
length does so by maintaining a constant percentage deviation from
the regression line (Klauber, 1941). The standard body length was
arbitrarily chosen as 400 mm. In determining the standard body
size, the writer was guided by the known biology of Amphiuma so
that the chosen body length would be within the adult (sexually
mature) body length range of the individuals of each of the six
series. The females of the Audubon Park population become sexually
mature when they attain a body length of 330 mm but the individuals
of the Audubon Park population do not attain a size as large as that
of Amphiuma collected from the surrounding areas (Cagle, 1948).
The 400 mm body length should include the sexually mature individ-
uals from all geographical series.

The body length range for the restricted number of specimens
was chosen to include the greatest number of individuals, the limit-
ing factor being the exclusion of individiuals less than 249 mm in
body length or greater than 600 mm in body length (Tables 3-6).
The statistical accuracy sacrificed by limiting the study to fewer
specimens (by taking a narrow range) is more than compensated for
by the elimination of possible errors in translating the specimens

No. 12 Hill: Taxonomic Status of Amphiuma 205

over too great a distance along their regression lines (Klauber, 1938).

The actual measurements of the body parts of each individual
within the restricted ranges were converted to the measurement they
would probably have at 400 mm body length by substituting the
actual measurements in a standardization formula (Klauber, 1941).

A survey of the available material indicated that a moderate number
of individuals (all sizes included) were available in Series 1 through
4, but the number in the restricted size range of Series 5 and 6 was
very small. The range could not be widened because the millimeter
spread on either side of the mid-point could not exceed 200 mm
(Klauber, 1938). Klauber has used as few as five individuals in his
snake studies.

In order to determine the standardized body parts for each individ-
ual of each series, the individuals from the restricted ranges of Series
1 through 6 were standardized by substituting the actual length of
the part and the body length in a standardization formula. The
estimated head length that the Amphiuma would have at the standard
body length of 400 mm is 34.33 mm. Standardized lengths of the
body parts, computed for the restricted number of specimens, were
gathered into an array (separate for each geographical series) and
their statistics computed (Tables 3-6).

Graphical relationships — Bar graphs were constructed to show the
relationship of the range, standardized mean, standard error of the
mean and the standard deviation of the four body parts for each of
the six series. The horizontal axis was calibrated to include the
range of the standardized measurements. For each series (geograph-
ical locality) the range of variation is shown by a horizontal line and
the mean (M) by a short vertical line. The blackened part of each
bar comprises two standard errors of the mean (2q-m) on either side
of M. One-half of each black bar plus the white bar at either end
outline one standard deviation (q-) on either side of M. Standard
deviation indicates dispersion and two standard errors of the mean
indicates reliability (Hubbs and Hubbs, 1953).

Results

Number of toes and coloration — The division of the 537 individ-
uals into the six geographical series shows the Audubon Park in-
dividuals (Series 2) to be the most numerically representative of
any of the series. Series 6, 5, 4, 1, and 3 follow, respectively, in
decreasing numbers. The division of the individuals into size groups
of 50 mm to determine the adequacy of the various age groups of
each series, yielded the following information: of the three-toed
groups, Series 1 was not represented in the size groups of less than
50 mm, of 250-300 mm, and of 350-400 mm; Series 2 was not repre-
sented in the size groups of 100-150 mm and 550-600 mm; Series 3
was not represented in the size groups of less than 50 mm, 100-150
mm, 200-250 mm, and 550-600 mm; Series 4 was not represented
in the size group of less than 50 mm. The groups not represented

206

Tulane Studies in Zoology

Vol. 1

in the two-toed populations by Series 5 are size groups of less than
50 mm, 300-350 mm, 450-500 mm, 500-550 mm, 550-600 mm; by
Series 6, 300-350 mm, 450-500 mm, and 500-550 mm (figures 6
and 7).

An inspection of the map showing the geographical distribution
of the individuals examined in this research indicates that the two-
toed and three-toed individuals occur together in the region where
the alluvial valley of the Mississippi River meets the East Gulf
Coastal Plain (figure 1). Within this region the proximity of the
three-toed Amphiuma to the two-toed Amphiuma varies. A col-
lection made in Amite, Louisiana, June 8, 1948, consisted of 42
Amphiuma taken from the same mud hole; of these, forty-one in-
dividuals had two toes on all four limbs and one individual had three
toes on all four limbs. There is reason to believe that these individ-
uals are from the same brood. All except one individual had a body
length of approximately 214 mm. In the same region individuals
with two toes on all four limbs and individuals with three toes on
all four limbs have been collected together, either in the same drain-
age ditches, in adjacent drainage ditches or within a few miles (less

if>

•

Figure 6. The distribution of body lengths of 290 specimens : Audu-
bon Park (cross-hatched), west of the Mississippi River (clear),
eastern alluvial valley (shaded), and Series 4 (stippled). The total
number of individuals composing1 each 50 mm size group is given at
the top of each bar of the histogram.

No. 12

Hill: Taxonomic Status of Amphiuma

207

o
o

IO
IO

Figure 7. The distribution of body lengths of 83 specimens from
Series 5 (shaded) and 163 specimens from the Atlantic coastal states
(cross-hatched). The number of individuals comprising each 50 mm
size group is given at the top of each bar of the histogram.

208 Tulane Studies in Zoology Vol. 1

than five) of each other.

Three individuals with both two-toed and three-toed limbs on the
same individual have been collected within this same geographical
area. One of these was collected in Mandeville, Louisiana, in August,
1924, and the other two individuals were collected three miles north
of the town of Pearl River, Louisiana. Both of these individuals
were collected within the critical area (Series 4 and 5). The one
from Mandeville (TU 2207) has a body length of 235.5 mm. The
ones collected north of Pearl River in January, 1949 (TU 4777),
and January, 1950 (TU 11414), have body lengths of 356 mm and
433 mm respectively.

Courtship activity between a two-toed and a three-toed Amphiuma
has never been reported. The males of the Audubon Park series are
reproductively active from mid-January to mid-May (Cagle, 1948).
It is possible that in the region where the alluvial valley of the
Mississippi River meets the East Gulf Coastal Plain, the two-toed and
three-toed Amphiuma are reproductively active at the same time.
The reason for the foregoing statement is that a two-toed male
Amphiuma collected one mile north of Covington, Louisiana, March
6, 1950, was in a reproductive condition. The cloaca of this male
was red, swollen and partially everted. The animal had more than
ten well defined teeth prints and cuts on its body. These teeth marks
and cuts may indicate fighting between the males during the repro-
ductive season (Cagle, 1948).

All individuals examined could be placed into one of the two
groups on the basis of number of toes except the three peculiar
individuals, or 0.56 percent of the population. The toe formula for
the individual from Mandeville is 2/2 3/3; for the two Pearl River
specimens, the formulae are 2/2 3/2 and 3/2 2/3. All individuals
of the six geographical series can be definitely separated into two-
toed groups and three-toed groups on the basis of color, with the
exception of the same three individuals which cannot be separated
into one of two groups on the basis of the number of toes.

Within the three-toed group there are various degrees of ventral
pigmentation, but the overall coloration is never so dark as that of
the two-toed individuals. Whether this variation in the ventral
coloration within the three-toed group is due to the effects of preser-
vation, ecology, or inheritance has not been determined. The per-
centage of individuals of two of the three-toed series with ventral
pigmentation is as follows: 7.8 percent for the Audubon Park in-
dividuals and 19.1 percent for those examined in Series 4. The
individuals from the eastern alluvial valley usually have a mottled
ventral surface.

Characteristically, the three-toed individuals have a black throat
patch. The two-toed individual has no throat patch. Two of the
three individuals, which cannot be separated into either the two-toed
or the three-toed groups on the basis of a consistant number of toes
and coloration, do not have a characteristic black throat patch, but

No. 12

Hill: Taxonomic Status of Amphiuma

209

one of the individuals does have an indication of a throat patch.

Body proportions — Although the ontogenetic spread is compara-
tively satisfactory for the six series, the frequencies of all size groups
should be increased, with the possible exceptions of the eastern two-
toed juveniles (50-100 mm) and the Audubon Park adults (300-
400 mm).

The correlation coefficients (r) are all high except that for fore-
limb length to body length of Series 3 (0.595). No explanation
is attempted for this low correlation coefficient.

The values for standard deviation (or) are calculated on the as-
sumption of normal distribution which is not likely to be realized
in small samples. There are reasons for distrusting the reliability of
the difference that is indicated by the graphical method when the
samples include fewer than thirty specimens. Consequently, the
limitation of the graphs (figures 8-11) is realized. The graphical
analysis indicates why a broad overlap of the dark bars indicates low
reliability of the observed difference between two samples and why
any considerable separation of these bars indictates a high reliability.
The plotting of one standard deviation on either side of the mean
indicates an 84 percent separation.

Figure 8. Bar graphs showing the relationship of the range, mean,
standard deviation and standard error of the mean for head length
for each of the six series. For each locality the range of variation is
shown by a horizontal line; the mean (M), by a short vertical line.
The blackened part of each bar comprises 2 standard errors of the
mean (2 crM) on either side of M. One-half of each bar plus the white
bar at either end outline 1 standard deviation ( o~) on either side of M.

210

Tulane Studies in Zoology

Vol. 1

3
4

•

.,_. 1

mm ■ i

^^ 1
1

. 1

*\ '

^^^n

^^^

8 9 10 II 12 13
SERIES FORE-LIMB LENGTH, mm.

Figure 9. Bar graphs showing the relationship of the range, mean,
standard deviation and standard error of the mean for fore-limb
length. Arrangement as in fig. 8.

Discussions and Conclusions

Number of toes and coloration — The separation of Amphiuma into
three groups on the basis of number of toes and coloration gives rise
to the question as to whether or not the smallest of the three groups,
comprising 0.56 percent of the population, is an intergrade group.
If the coloration of these three individuals is due to either ecology
or to methods of preserving the animals, and if the variation in the
number of toes is due to either mutilation or to the inheritance of
the monodactyly or Polydactyly type, then Amphiuma can be com-
pletely separated into two groups, indicating no intergradation.

Since other individuals occurring within the same ecological area do
not have the same coloration as these three individuals, indicates that
ecology per se is not the causative factor in producing this coloration,
unless the ecological factor producing it is extremely localized. The
latter assumption is unlikely; for, if Amphiuma is so susceptible to
these extremely localized conditions, a greater degree of natural vari-
ation would be expected among the remaining individuals than has
been found. It is interesting that the type of coloration shown by
these three individuals, which may be a blending resulting from the
interbreeding of a bicolored individual with a unicolored individual,
is found only in the region where the ranges of the two-toed and the
three-toed individuals meet and overlap.

The fact that these three animals have no apparent scar tissue and
that the measurements of the left limbs are the same as those of
their corresponding right limbs constitutes evidence against the oc-
currence of mutilation. It is true that monodactyly and Polydactyly

No. 12

Hill: Taxonomic Status of Amphiuma

211

Figure 10. Bar graphs showing the relationship of the range, mean,
standard deviation and standard error of the mean for hind-limb
length. Arrangement as in fig. 8.

100 115 130 145 160

SERIES TAIL LENGTH, mm.

175

190

Figure 11. Bar graphs showing the relationship of the range, mean,
standard deviation and standard error of the mean for tail length.
Arrangement as in fig. 8.

212 Tulane Studies in Zoology Vol- 1

occur in Ampbiuma, but it is not evident that either one of these
conditions expresses itself in any of the three individuals. Poly-
dactyly has been observed only in individuals in the three-toed series,
which series were designated by number of toes, coloration (general
body color and the presence of throat patch), and geographical
location. TU 4777 would have to be considered a three-toed individ-
ual because of the presence of the throat patch. The toe formula
2/2 2/3 must then be a result of inheritance (didactyly) as there
is no evidence of mutilation. TU 2207 and TU 11414 because of
the absence of a throat patch would be considered two-toed individ-
uals. The toe formulae, 2/2 3/3 and 3/2 2/3, respectively, for
TU 2207 and TU 11414 must be explained also by inheritance
(Polydactyly); however, Polydactyly has not been observed in two-
toed individuals (concerning geographical ranges) in this collection.
In spite of the assumption of the presence of Polydactyly, and the
fact that the throat patch separates the three individuals into two
groups, the uniform coloration of the three individuals and the almost
identical general coloration of two of the individuals, TU 4777 with
the throat patch and TU 11414 without the throat patch, prevents
these three individuals from being separated into one of two groups.

Basically, the taxonomic position of the Mid-Gulf Coast Ampbiuma
resolves itself into the taxonomic position of these three individuals;
that is, whether or not the three can be separated into one of two
groups, a two-toed or a three-toed group, or whether all three should
be placed into a third group, an intergrade group. The question is
whether these three intermediates are true subspecies or hybrids.

With the exception of three specimens, all two-toed specimens
have one toe pattern and all three-toed specimens have another. Since
the two tables (1 and 2) are generally concordant and the forms
(consistent with respect to two different characters, number of toes
and pattern) are also sympatric in southeastern Louisiana (at least),
and, in some cases, in the same pond, we are probably dealing with
two separate species. It is difficult to conceive of two characters as
different as number of toes and pattern that would result from a
single genie element. If we were dealing with true subspecies and
an area of intergradation, we should expect to find an almost com-
plete mixture of characters in the intergrading area; that is, two-
toed individuals with all variations in pattern and three-toed individ-
uals with the same diversity. Intergradation is indicated in only
three specimens out of 131 from the critical area (Series 4 and 5).

On the basis of this investigation the writer considers the three
individuals to be hybrids. If there were a high proportion of mixed
individuals in the critical area, the case for intergradation and a sub-
specific relationship would be greatly strengthened, particularly if
the characters typifying the two groups were well jumbled in the
intermediates. More individuals need to be collected from the critical
area (Series 4 and 5).

Body proportions — The indication of species separation is further

No. 12 Hill: Taxonomic Status of Amphiuma 213

reinforced by the fact that three-toed individuals, as a group, differ
from the two-toed in proportionate limb length, and that the differ-
ence exceeds the within-group divergence (Tables 3 and 4). This
is probably one of the most valuable parts of the investigation. It
is not as important to note that head and tail portionalities are similar
as to note that the latter reinforce and substantiate the fundamental
toe and color pattern differences.

Summary

Amphhima has been separated into two species, Amphiuma means
and Amphiuma tridactylum, by Baker on the basis of number of toes,
color and the ratios of total length to body length and body length
to limb lengths. In deriving the mean ratios, Baker did not recognize
the influence of ontogenetic variation, allometric growth or sexual
dimorphism.

This writer's research was concerned with the evaluation of the
three assumptions above and an attempt to determine the real dif-
ferences between the two-toed groups and the three-toed groups.

Heretofore, Amphiuma has been separated into a maximum of
three groups to be compared with each other. Goin divided Am-
phiuma into a three-toed group, a two-toed group and an intergrade
group and compared these groups. The author divided 537 Am-
phiuma into six series on the basis of geographical distribution.
Series 1 is composed of individuals collected west of the Mississippi
River; Series 2 consists of individuals from Audubon Park; Series 3
includes those individuals within the alluvial valley east of the
Mississippi River; Series 4 (three-toed) and Series 5 (two-toed)
includes those Amphiuma within the region where the alluvial valley
of the Mississippi River meets the East Gulf Coastal Plain; Series 6
is composed of individuals from the Atlantic coastal states.

The reason for this division of Amphiuma into series was to deter-
mine the range of variation within the three-toed series and within
the two-toed series so that the differences between two-toed and
three-toed Amphiuma could be interpreted more accurately. The
Audubon Park series was separated from all other series because it
is geographically isolated and the individuals of this series do not
reach a size as large as those from the surrounding areas. Since it
was not known whether or not the three-toed Amphiuma within the
eastern alluvial valley of the Mississippi River were genetically
similar to those west of the Mississippi, these two series were sepa-
rated. Series 5 was separated from Series 6 in order to determine
if there was any difference between these two two-toed groups and
to determine if there was a closer relationship between Series 5 and
Series 6 than between Series 4 and Series 5.

The characters used in determining the differences and/or simi-
larities between the series are number of toes, coloration, body pro-
portions of head length to body length, fore-limb length to body
length, hind-limb length to body length and tail length to body length.
The two-toed and the three-toed Amphiuma cannot be completely

214 Tulane Studies in Zoology Vol. 1

separated into two groups by any one of the above characters. The
best separation is obtained on the basis of number of toes and throat
coloration.

Intergradation is indicated in only three specimens out of 131
from the critical area (Series 4 and 5). On the basis of this investi-
gation the three individuals are considered hybrids. The indication
of species separation is further reinforced by the fact that three-toed
individuals, as a group, differ from the two-toed in proportionate limb
length, and that the difference exceeds the within-group divergence.

Acknowledgements
I am indebted to Dr. Fred R. Cagle of the Department of Zoology,
Tulane University, for suggesting this problem and for many helpful
suggestions during the course of this study and the preparation of
this manuscript. I am indebted to Drs. E. S. Hathaway and G. H.
Penn for their critical analysis. For the loan of specimens, I wish
to thank Mr. Charles M. Bogert of the American Museum of Natural
History, Dr. Doris M. Cochran of the United States National Museum,
Mr. Richard Etheridge of Houston, Texas, and Dr. Arnold B. Grobman
of the University of Florida. Mr. Allan H. Chaney and other graduate
students at Tulane University were helpful in collecting additional
field samples after this research was begun.

References Cited

Baker, C. L. 1945. The natural history and morphology of amphi-
umae. Jour. Term. Acad. Sci., 20: 55-91.

_. 1947. The species of amphiumae. Jour. Tenn. Acad.

Sci., 22: 9-21.

Brimley, C. S. 1909. Some notes on the zoology of Lake Ellis, Craven
County, North Carolina, with special reference to herpetology.
Proc. Biol. Soc. Wash., 22: 129-137.

Cagle, Fred R. 1948. Observations on a population of the salamander,
Amphiuma tridactylum Cuvier. Ecology, 29: 479-491.

Cope, E. D. 1886. Synonymic list of the North American species of
Bufo and Rana, with descriptions of some new species of Batra-
chia, from specimens in the National Museum. Proc. Amer. Phil.
Soc, 23: 514-526.

Cuvier, M. Le Bon 1827. Sur le genre de reptiles batraciens, Nomme
Amphiuma, et sur une nouvelle espece de ce genre (Amphiuma
tridactylum). Mem. Mus. Hist. Nat. Paris, 14: 7.

Fisher, R. A. 1938. Statistical Methods for Research Workers,
(7th ed.). Oliver and Boyd, London, pp. 1-356.

Fitzinger, Leopoldo 1843. Systema Reptilium, Fasciculus Primus
Ambyglossae, 106, IX p.

Goin, C. J. 1938. The status of Amphiuma tridactylum Cuvier. Her-
petologica, 1 : 127-130.

Hubbs, Carl L. and Clark Hubbs 1953. An improved graphical anal-
ysis and comparison of series of samples. Systematic Zoology,
2: 49-56, 92.

No. 12 Hill: Taxonomic Status of dmphiuma 215

Klauber, L. M. 1938. A statistical study of the rattlesnakes : V. Head
dimensions. Occ. Pap. San Diego Soc. Nat. Hist., No. 4: 1-53.

1941. Four papers on the application of statistical

methods to herpetological problems. Bull. Zool. Soc. San Diego,
No. 17: 5-71.

1943. Tail length differences in snakes with notes

on sexual dimorphism and the coefficient of divergence. Bull.
Zool. Soc. San Diego, No. 18 : 1-60.

Peatman, John G. 1947. Descriptive and Sampling Statistics. Har-
per and Brothers, New York, pp. 1-577.

Ryder, John A. 1880. Morphological notes on the limbs of amphi-
umae, as indicating a possible synonomy of the supposed genera.
Proc. Acad. Nat. Sci. Phila., 1879: 14-15.

Stejneger, Leonhard and Thomas Barbour 1943. Check List of
North American Amphibians and Reptiles (5th ed.). Harvard
University Press, Cambridge, pp. 3-4.

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