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FIELDIANA . ZOOLOGY

Published by
CHICAGO NATURAL HISTORY MUSEUM

Volume 34 NOVEMBER 13, 1956 No. 41

GEOGRAPHIC VARIATION IN THE
CHICKEN TURTLE

Deirochelys reticularia Latreille

ALBERT SCHWARTZ

DEPARTMENT OF BIOLOGY, ALBRIGHT COLLEGE, READING, PENNSYLVANIA

The chicken turtle, Deirochelys reticularia, was described as
Testudo reticularia by Latreille in 1801. Almost simultaneously, the
same turtle was described by Daudin as Testudo reticulata. Both
descriptions were based upon data gathered by Bosc in North
America (1798-1800). During his visit to the United States, Bosc
spent some time at Charleston, visiting with his countryman,
Andre" Michaux, and presumably his collection of the original
specimen and his observations on the chicken turtle were made
at that time. The provenance of this specimen, in the custom of
that period of zoological taxonomy, was stated merely as "Caro-
lina"; the Michaux Garden was located about twelve miles north
of Charleston, and the type specimen may well have come from this
vicinity. Harper (1940, p. 711) and Schmidt (1953, p. 104) restricted
the type locality to Charleston, South Carolina, and this restriction
is appropriate.

The question of the priority of Testudo reticularia Latreille over
Testudo reticulata Daudin has been ably and thoroughly discussed
by Harper (op. cit., pp. 710-711). According to this author, the
Latreille name antedates the Daudin name by several mc.iths and
thus is the valid name for the chicken turtle of the southern United
States.

Agassiz first recognized the distinctive characters which differ-
entiate Deirochelys from other emydid genera. Until that time, the
chicken turtle had been placed in three genera (Testudo, Emys,
Terrapene) and subsequently has been considered as belonging to
the genera Clemmys and Chrysemys. Agassiz separated Deirochelys
from the remaining Emydidae on the basis of the long neck and the
mode of articulation of the neck vertebrae. Even more striking, as

NO ' 805 461 THEU3WWOFTK:

NAT.

MOV 1866 HIST.

462 FIELDIANA: ZOOLOGY, VOLUME 34

Carr (1952, p. 316) has pointed out, is the slimness of the free upper
ends of the ribs, which can thus accommodate the large retractor
muscles of the neck. From all other North American emydids
except Emys and Clemmys marmorata, Deirochelys differs in that the
first central lamina is in contact with five rather than three marginal
laminae. Williams (1950) did not examine specimens of Deirochelys
in his study of the cervical articulation of turtles, and skeletons
have not been available to me. Such study should prove valuable
and interesting, because of the elongate neck and presumed con-
comitant muscular and skeletal modifications of Deirochelys.

No one has made mention of geographic variation in Deirochelys
reticularia, and indeed little seems to be known regarding the
chicken turtle in general. However, Carr (op. cit., p. 318) noted
that a hatchling from southern Louisiana possessed a "dark linear
pattern following the seams of the plastron." This character, often
somewhat modified through age, occurs consistently in populations
west of the Mississippi River and distinguishes these western turtles
from those of the eastern United States. Coloration and carapace
shape serve also to differentiate populations of Deirochelys on the
Florida peninsula and the adjacent continental mass, so that three
subspecies are herein recognized, two of which are described as new.

Three hundred and twenty-five specimens have been examined,
with the following distributions by states: Oklahoma, 3; Arkansas, 8;
Texas, 42; Louisiana, 57; Mississippi, 7; Alabama, 5; Georgia, 52;
Florida, 113; South Carolina, 32; North Carolina, 6. Most of these
are entire specimens, but a few are dry shells or preserved heads
only. The chicken turtles from the following collections have been
examined. I wish to express my thanks to the respective curators of
these collections and to private individuals for allowing me to borrow
study material in their care: American Museum of Natural History,
Charles M. Bogert (AMNH); Arkansas Polytechnic College, Allan
H. Chaney (APC) ; Chicago Academy of Sciences, Howard K. Gloyd
(CAS); Charleston Museum (ChM); Carnegie Museum, Neil D.
Richmond (CM); Chicago Natural History Museum, Robert F.
Inger (CNHM); Cornell University, Edward C. Raney (CU);
Museum of Comparative Zoology, Arthur Loveridge (MCZ);
Strecker Museum, Bryce C. Brown (SM); Texas Cooperative Wild-
life Museum, W. B. Davis (TCWM); Texas Natural History Col-
lection, University of Texas, W. F. Blair (TNHC); Tulane Uni-
versity, Fred R. Cagle (TU) ; United States National Museum, Doris
M. Cochran (USNM); University of Florida, Archie F. Carr, John
W. Crenshaw and Duke Wilder (UF); University of Georgia,

' SCHWARTZ: CHICKEN TURTLE 463

Bernard Martof; University of Illinois, Museum of Natural History,
Hobart M. Smith (UIMNH); University of Kansas, Edward R.
Taylor (UK); University of South Carolina, Harry W. Freeman;
University of Michigan, Museum of Zoology, Norman E. Hartweg
and William E. Duellman (UMMZ); University of Oklahoma, Mu-
seum of Zoology, Charles C. Carpenter (UOMZ). Specimens from
private collections have been lent by Bryce C. Brown (BCBC),
John W. Crenshaw, Richard E. Etheridge, Edmund R. Cuthbert,
and Robert L. Humphries. L. Neil Bell, Larry H. Ogren, Dennis
R. Paulson, and Donald W. Tinkle have supplied me with living
specimens of turtles from Florida, Texas, and Louisiana, and Adrian
A. Armour, Julian R. Harrison III, John A. Quinby, Daniel R.
Stanland and others have collected specimens for me in South
Carolina. The specimens in the University of South Carolina were
collected during a survey of the herpetofauna of the Savannah
River Plant, sponsored by the Atomic Energy Commission. I wish
to thank Doris M. Cochran, Roger Conant, James A. Peters, and
Arthur Loveridge for assistance in checking references in certain
literature. The plates are the work of Isabelle Hunt Conant, and
I wish to thank her for her faithful reproductions of the original
specimens.

Carr's (1952, p. 49) method was used in taking shell measure-
ments; measurements were taken with vernier calipers when possible,
and larger individuals were measured with dividers for all measure-
ments except depth, for which calipers were employed. All measure-
ments are in millimeters. The following abbreviations are used:
CL, greatest length of carapace; PL, greatest length of plastron;
C3/4, width of carapace at seam between marginals 3 and 4; C5/6,
width of carapace at seam between marginals 5 and 6; C8/9, width
of carapace at seam between marginals 8 and 9; CW, greatest width
of carapace; D, greatest depth of shell. The head width was taken
at the level of the anterior border of the tympanum. All capital-
ized color names are from Ridgway (1912).

In the subspecies discussions, specimens have been segregated
into three categories: (1) Juveniles (individuals from hatchling size
to about 79 mm.). This group is generally characterized by the
retention of the primary or hatchling carapace laminae surrounded
by a smooth or lightly striated zone of new growth. (2) Subadults
(individuals with finely striated laminae which have not reached
adult size. The size range is from about 68 mm. to 141 mm.). The
placing of a single specimen in either the juvenile or the subadult
category depends upon the appearance of the carapace laminae,

464 FIELDIANA: ZOOLOGY, VOLUME 34

and this accounts for the overlap in carapace measurements of these
two groups. (3) Adults (males with carapace lengths in excess of
96 mm. and females with carapace lengths in excess of about 155
mm. except in the subspecies miaria where females are considered
adult at lengths of 139 mm. and over). The subadult and adult
categories overlap because large subadult females are larger than
small adult males.

Latreille (1801, p. 124) stated that the specimen of Testudo
reticularia available to him measured 7 inches in carapace length,
4:^2 inches in width, and 3 inches in depth. These dimensions agree
with those of a female from Aiken County, South Carolina, and,
since male Deirochelys in South Carolina do not reach this size, the
original Bosc specimen was almost certainly an adult female.
Latreille (op. cit., p. 126) said that the plastron was yellow, and this
affirms the fact that the original specimen did not come from
Florida. M. Jean Guibe" of the Museum National d'Histoire Natur-
elle has informed me (letter, December 13, 1954) that the type
specimen of Testudo reticularia Latreille "doit etre consideYe" comme
perdu." With the description of two new subspecies of this turtle
in the present paper, it seemed suitable to designate both a neotype
and a neoallotype from the vicinity of Charleston, South Carolina,
and, since the type specimen was almost certainly a female, a female
has been selected as the neotype.

Genus Deirochelys Agassiz

Deirochelys Agassiz, 1857, Contr. Nat. Hist. U. S., 1 : 441.

Dirochelys Baur, 1889, Amer. Nat., 22: 1099-1100.

Dierochelys Loding, 1922, Alabama Mus. Nat. Hist., mus. paper no. 5, p. 45.

Diagnosis. An emydid turtle with an oval or cuneiform cara-
pace outline; laminae of carapace dark, striated, and with a reticu-
lum of yellowish lines and usually a light border; plastron unhinged
and immovable, yellow or orange in color, sometimes with a seam-
following dark plastral pattern; bridge with or without a dumbbell-
shaped figure; shell deep or flattened; marginal laminae 25, often with
a dark blotch on the ventral surface; four pairs of lateral laminae;
five central laminae; neck elongate and about equal to length of
carapace; skin black or brown with lines and markings yellowish or
greenish; a wide band of yellow on anterior surface of forelimb;
a series of vertical, alternating black and yellow bars between tail
and hind limbs on rump; free ends of ribs very slim for accommoda-
tion of large retractor muscles of neck; first central lamina in con-

SCHWARTZ: CHICKEN TURTLE 465

tact with five marginal laminae (precentral, marginals 1 and 2 on
each side).

Deirochelys reticularia reticularia Latreille

Testudo reticularia Latreille, 1801, in Sonnini and Latreille, Hist. Nat. Rept.,
1: 124, pi. 6, fig. 1.

Testudo reticulata Daudin, 1801, Hist. Nat. Rept., 2: 144, pi. 21, fig. 3; Le
Conte, 1830, Ann. Lye. Nat. Hist. New York, 3: 103.

Emys reticulata Schweigger, 1812, Konigsberg. Arch. Naturwiss. Math., 1
(3), pp. 300, 346, 1 (4), p. 425; Schweigger, 1814, Prod. Mon. Chelon.,
pt. 2, p. 31; Merrem, 1820, Tent. Syst. Amphib., p. 26; Dumeril and
Bibron, 1835, Erpet. Gen., 2: 291; Holbrook, 1838, N. Amer. Herp., 1st
ed., 2: 41, pi. 7; Holbrook, 1842, N. Amer. Herp., 2nd ed., 1: 59, pi. 7;
Holbrook, 1848, in White, Statistics of Georgia, Fauna and Flora, p. 13;
Le Conte, 1854, Proc. Acad. Nat. Sci. Philadelphia, 7: 189.

Emys reticularia Gray, 1844, Cat. Turtles Brit. Mus., p. 25; Gray, 1855, Cat.
Shield Reptiles Brit. Mus., p. 27.

Terrapene reticulata Bonaparte, 1830, Osserv. Seg. Ed. Regno Anim., p. 155;
Bonaparte, 1830, Ann. Stor. Nat. (Bologna), 4 (3), p. 369.

Deirochelys reticularia Gray, 1870, Suppl. Cat. Shield Reptiles Brit. Mus.,
pt. 1, p. 39; Stejneger and Barbour, 1917, Check List of N. Amer. Am-
phibians and Reptiles, p. 121 (part); Carrington, 1929, Copeia, 172: 82;
de Sola and Abrams, 1933, Copeia, 1: 11; Pope, 1939, Turtles U. S. and
Canada, pp. 234-235 (part); Brimley, 1939-43, Reptiles and Amphibians
N. Car., Carolina Tips, Installment 31, second unnumbered page (part);
Carr, 1940, Univ. Fla. Publ., Biol. Sci. Ser., 3 (1), p. 105 (part); Jopson,
1940, Herpet., 2 (2), p. 43; Ditmars, 1946, Reptiles N. Amer., pp. 418-419
(part), pi. 120, lower fig.; Smith and Buechner, 1947, Bull. Chicago
Acad. Sci., 8 (1), pp. 1-16; Neill, 1948, Herpet., 4 (3), p. 114; Brown,
1950, Check List of Reptiles and Amphibians, Texas, p. 234 (part); Carr,
1952, Handbook of Turtles, pp. 316-319 (part); Chermock, 1952, Geol.
Surv. Alabama, mus. paper no. 33, p. 51; Schmidt, 1953, Check List of
N. Amer. Amphibians and Reptiles, p. 104 (part); Carr and Goin, 1955,
Reptiles, Amphibians, and Fresh-water fishes of Fla., p. 246 (part);
Freeman, 1955, Univ. S. Car. Publ. Biol., ser. 3, 1 (1), pp. 241-243;
Mertens and Wermuth, 1955, Zool. Jahrb. (Syst.), 83, (5), pp. 348-349
(part); Smith and List, 1955, Amer. Mid. Nat., 53 (1), p. 125.

Deirochelys reticulata Agassiz, 1857, Contr. Nat. Hist. U. S., 1: 441, 2, pi. 1,
figs. 14-16, pi. 7, figs. 17-19; Siebenrock, 1909, Zool. Jahrb., suppl. vol.
10 (3), p. 485 (part); Deckert, 1918, Copeia, 54: 31.

Dirochelys reticularia Baur, 1889, Amer. Nat., 22: 1099-1100.

Clemmys reticularia Strauch, 1862, Mem. Acad. Sci. St. Petersburg, (7), 5
(7), p. 32; Strauch, 1865, Mem. Acad. Sci. St. Petersburg, 8 (13), p. 78.

Chrysemys reticulata Cope, 1875, Bull. U. S. Nat. Mus., 1 : 53; Davis and Rice,
1883, Bull. Illinois State Lab. Nat. Hist., 1 (5), p. 56; True, 1883, Bull.
U. S. Nat. Mus., 24: 35 (part); True, 1884, Fish. Indust. U. S., sec. 1,
p. 157 (part); Boulenger, 1889, Cat. Chel. Brit. Mus., p. 75 (part); Lind-

466 FIELDIANA: ZOOLOGY, VOLUME 34

holm, 1901, in Lampe, Cat. Kept. Samml. Naturhist. Mus., Wiesbaden,

p. 10.
Chrysemys reticulatus Wright and Funkhouser, 1915, Proc. Acad. Nat. Sci.

Philadelphia, 67: 117.
Dierochelys reticularia Loding, 1922, Alabama Mus. Nat. Hist., mus. paper

no. 5, pp. 45-46 (part); Haltom, 1931, Geol. Surv. Ala., Alabama Mus.

Nat. Hist., mus. paper no. 11, p. 137 (part).

Type locality. Carolina; by restriction, vicinity of Charleston,
Charleston County, South Carolina.

Neotype. ChM 54.48.1, adult female, taken 9 miles northwest
of Charleston, Charleston County, South Carolina, on April 1, 1954,
by T. A. Beckett.

Neoallotype. ChM 52.140.5, adult male, taken 2.1 miles north,
thence 1.2 miles west of Cainhoy, Berkeley County, South Carolina,
on December 20, 1952, by Julian Harrison III and John A. Quinby.

Specimens examined. NORTH CAROLINA: Craven Co., New Bern,
2; Carteret Co., Barker's Island, Willow Pond, 1; New Hanover Co.,
Wrightsville, 1, Wilmington, 1; Brunswick Co., Ft. Caswell Beach, 1.
SOUTH CAROLINA: Williamsburg Co., Kingstree, 1, Lanes, 1; Claren-
don Co., Santee National Wildlife Refuge, Lake Marion, 1 (not
mapped); Georgetown Co., Kinloch Plantation, 12 miles south of
Georgetown, 1; Berkeley Co., Lake Moultrie, near Pinopolis, 1
(not mapped), Medway Plantation, Back River, 1, 0.9 miles north
of Cainhoy, 1 (not mapped), 6 miles north of Cainhoy, 1, 2.1 miles
north, thence 1.2 miles west of Cainhoy, 8; Charleston Co., Drayton
Station, 1; Dorchester Co., Summerville, 1; Aiken Co., 3 miles
northeast of Ellenton, 3, pond at point where State Highway 19
crosses Tim's Prong of Upper Three Runs Creek, 1, northwest edge
of Ellenton, 1, Savannah River area, 1; Barnwell Co., 1 mile west
of Snelling on South Carolina Route 64, thence 1 mile south of South
Carolina Route 64, 4; Jasper Co., Great Swamp, tributary of New
River, 1; Beaufort Co., Bluffton, 1. GEORGIA: Wayne Co., near
Liberty County line, 1; Lee Co., Usry Plantation, 1; Worth Co.,
9.7 miles north of Sylvester, 1; Baker Co., Mimsville, 12, 4 miles
northeast of Emory Field Station, 6, 2.0 miles east of Miller County
line, 1, 0.1 miles south of Dougherty-Baker County line on U. S.
Highway 91, 1, 100 yards south of Dougherty-Baker County line, 1,
Thompson Pond, ca. 5 miles northwest of Newton, 1, no other
locality, 2; Grady Co., Beachton, Birdsong Plantation, 1; Thomas
Co., 8 miles south of Thomasville, 1, no other locality, 1; Clinch
Co., vicinity of Stockton, 1, vicinity of Homerville, 2; Lowndes Co.,
Valdosta, 1; Ware Co., 9 miles southeast of Waycross, 1, Okefinokee

467

468 FIELDIANA: ZOOLOGY, VOLUME 34

Swamp, 2, Okefinokee Swamp, Chesser's Island, 1, Okefinokee
Swamp, Honey Island, 1; Charlton Co., Camp Cornelia, 1, Folkston,

1, 1 mile south of Trader's Hill, 1. FLORIDA: Escambia Co., Pensa-
cola, 2; Washington Co., near Chipley, 1; Bay Co., 4 miles west of
Panama City, 1; Calhoun Co., 6 miles south of Blountstown, 4,
Chipola River, 4 miles south of Scott's Ferry, 1; Gulf Co., 10 miles
west of Wewahitchka, 1; Gadsden Co., 13 miles northwest of
Tallahassee, 1; Leon Co., 2 miles southwest of Tallahassee, 1;
Franklin Co., east of Eastpoint, 1, Mexico Beach, 1; Jefferson Co.,
Yi mile northwest of Lament, Highway 24, 1; Madison Co., 1 mile
southeast of Lamont, 1; Nassau Co., 1.7 miles south of Hilliard, 1.
ALABAMA: Greene Co., Tombigbee River, 2; Dallas Co., 1.4 miles
northeast of Orrville, 1; Mobile Co., Mobile, 1; Baldwin Co., 9 miles
north of Foley, 1. MISSISSIPPI: Stone Co., 1; Harrison Co., Biloxi,

2, 10 miles north of Biloxi, 1; Jackson Co., Moss Point, 1, 16 miles
north of Van Cleave, 1, 2.5 miles north of Hurley, 1. LOUISIANA:
Washington Parish, 11 miles west of Bogalusa, 1 (not mapped);
St. Tammany Parish, 8 miles from Pearl River, 3, Sun, 1; Tangi-
pahoa Parish, 5 miles east of Hammond, 1.

Geographic distribution. The Atlantic and Gulf coastal plains,
from east central North Carolina through South Carolina, Georgia,
northern and western Florida, Alabama, Mississippi, and the Florida
Parishes of Louisiana (see map, fig. 102). Intergrades between D. r.
reticularia and D. r. chrysea occur in north central Florida in Taylor,
Dixie, Levy, Gilchrist, and Clay counties.

Diagnosis. Deirochelys r. reticularia differs from D. r. chrysea
in the generally less bright coloration of the former, the much
narrower dorsal reticulation, which is greenish or brownish, the
narrow yellow edging of the carapace, and the sporadic occurrence
of a plastral spot at the juncture of the femoral and anal laminae.
The black spots on the ventral side of the marginals at the level of
the bridge are present in 72 per cent of the specimens examined;
comparison with specimens of D. r. chrysea shows that 43 per cent
of these specimens have marginal black spots at the level of the
bridge. A chi square test for these data indicates that this differ-
ence is significant.

The shape of the carapace distinguishes D. r. reticularia from
D. r. chrysea. The shell is ovate in shape, and not cuneiform and
expanded as in D. r. chrysea; there is seldom the constriction at the
level of marginals 5 and 6 that occurs in D. r. chrysea. D. r. reticu-
laria is flatter than D. r. chrysea. See Tables 1 and 2.

SCHWARTZ: CHICKEN TURTLE 469

D. r. reticularia differs from D. r. miaria in never having the
dendritic plastral pattern following the seams. When a plastral
pattern is present in D. r. reticularia, it is a poorly defined blotch,
with occasional antero-median extension, at the femoral-anal suture.
Also, D. r. reticularia is less flat than D. r. miaria.

Description of neotype. An adult female (fig. 103) with the
following measurements: CL, 199; PL, 182; C3/4, 120; C5/6, 144;
C8/9, 128; CW, 144; head width, 30.5; D, 83; width of posterior
lobe of plastron, 79.3; width of bridge, 63.2. Ratios: D/CW, 0.58;
C5/6/CL, 0.72, CW/CL, 0.72. The shell is ovate in outline when
viewed dorsally, and highly domed; the greatest width of the cara-
pace is at the level of marginal 7 and the greatest height at central 3.
The posterior marginals are moderately flaring, and the lateral
marginals almost vertical. The carapace is dark brown in color, and
the reticulations narrow (not more than 1.5 mm. in width). The
carapace is edged with yellow for its entire margin, but only nar-
rowly so. The plastron is yellow. The dumbbell-shaped figure on
the bridge is separated into two separate elongate hollow blotches
at the pectoral-abdominal suture. The ventral surfaces of marginals
4, 5, and 6 have solid black spots on the right side and marginals
4 and 5 have black spots on the left. These marginal spots are oval
and are restricted to a single marginal except for a slight overlap
of the pigment onto the next most posterior marginal.

The head skin is black and the yellowish lines are prominent.
Five lines enter the posterior margin of the eye, the most dorsal and
most ventral (orbital) lines being the widest and most prominent.
The median head line is joined unilaterally to the left canthal line
and extends posteriorly about 6 mm. behind the eye. Two para-
median head lines begin just posterior to the eye and extend posteri-
orly onto the neck. A post-ocular line begins at the corner of the
eyelids and extends posteriorly as a broadening stripe. Each of
these head lines is separated by three faint yellow lines. The dorsal
and lateral aspects of the neck are black and the lines only faintly
discernible. The chin has a horseshoe-shaped yellow figure with
the open end projecting posteriorly and extended as two broad
yellow stripes onto the neck. An equally broad yellow stripe begins
at the apex of the horseshoe and extends posteriorly on the mid-
ventral line of the neck. There is a yellow line on the upper beak
extending from the nares to the margin of the beak and continued
as a blotch on the lower beak, which is elsewhere striped with
yellow and black.

470 FIELDIANA: ZOOLOGY, VOLUME 34

The forelimbs have a wide (about 8 mm.) band of yellow across
the anterior aspect at the level of digits 3 and 4, and the ventral
edge of the forelimb has a yellow line. The webs between the digits
are yellow. Between the elbow and the carapace there are 9 yellow
lines on the black skin. The hind limbs are black dorsally and yellow
ventrally except for the palmar surfaces of the feet, which are black.
The webs are yellow. Between the knee and the vent there are
7 to 9 yellow vertical lines on the black skin. The tail is terete and
short (31 mm. from vent to tip of tail), and the vent lies immediately
posterior to the posterior margin of the plastron.

Description of neoallotype. An adult male (fig. 104) with the
following measurements: CL, 141; PL, 127; C3/4, 83; C5/6, 96;
C8/9, 92; CW, 101.4; head width, 24.0; D, 60; width of posterior
lobe of plastron, 58.2; width of bridge, 42.0. Ratios: D/CW, 0.59;
C5/6/CL, 0.68; CW/CL, 0.72. The shell is ovate in outline and
moderately high on the mid-dorsal line; the greatest width of the
carapace is at the level of marginal 7 and the greatest height at
central 3. The posterior marginals are moderately flaring, especially
marginals 7, 8, and 9, and the lateral marginals are almost vertical.
The carapace is dark brown in color, with narrow reticulations and
mid-dorsal line as in the female. The carapace is edged with yellow
but not widely or prominently so, except at the confluence of the
reticulum with the yellow marginal border. The plastron is yellow.
The bridge pattern is almost absent, the dumbbell-shaped figure
being restricted to a diffuse small blotch on the inguinal laminae.
The ventral surfaces of the marginals are immaculate except for
small dots on marginals 4 and 5 on the right side and 3 and 4 on
the left.

The head pattern resembles that of the neotype in most details
but differs in that the median head line is not attached to the canthal
line and the paramedian I and the post-ocular lines are joined
on each side by a transverse yellow line at the level of the anterior
third of the tympanum. There are three wide yellow bands on the
neck, but no horseshoe-shaped figure as in the neotype. The
coloration and pattern of the limbs are essentially those of the female
described above. The tail is stout, cylindrical, and tapering (26 mm.
from vent to tip of tail), and the vent is about 16 mm. posterior to
the rear margin of the plastron. The penis is extruded and is uni-
formly black except for the pale yellow proximal ventral surface.

In the following discussions, specimens of D. r. reticularia from
western and northern Florida and southern Georgia have not been
employed.

o

ta cS

471

472

FIELDIANA: ZOOLOGY, VOLUME 34

FIG. 104. Dorsal and ventral views of the neoallotype of Deirochelys r.
reticularia (ChM 52.140.5) from 2.1 miles north, thence 1.2 miles west of Cainhoy,
Berkeley County, South Carolina.

JUVENILES

The sixteen juvenile individuals available for study range in
carapace length between 29.7 and 55.3 mm. The ratio of CW to CL
varies between 0.85 and 0.97, with the smaller individuals having
the higher ratios; in very small individuals the shell is more nearly
circular in outline. With increasing growth, the shell becomes
longer and somewhat broader posteriorly in the region of marginals
6 and 7, but it seldom reaches the extremely cuneiform configuration
of the Floridian form. In general, the ground color of the carapace
is dull brownish to olive, and in the smallest individuals the reticu-
lum and mid-dorsal lines are yellow and obvious, but are not nearly

SCHWARTZ: CHICKEN TURTLE 473

so broad as in juvenile D. r. chrysea. The shell is margined with
yellow, but not to so prominent a degree as in the more southern
subspecies. The carapace is high mid-dorsally, and the ratio of
D to CW varies between 0.45 and 0.58. In the largest turtle here
considered juvenile, the reticulations are already fine and obscured
by the darker coloration of the carapace. One specimen (TU 6040)
has the only deformed carapace in the entire series of Deirochelys
examined; the centrals have grown to the detriment of the left
lateral laminae, which are compressed in an antero-posterior direc-
tion and less developed than the right laterals.

The plastron is immaculate and yellow in all but four individuals;
in these four specimens there is a small but well-defined spot at the
juncture of the femoral and anal laminae. The bridge may be
immaculate or may have a pattern. Two specimens have the bridge
without either the dumbbell-shaped blotch or the spots on the
marginals, while seven individuals have both the figure and the
marginal spots. The remaining seven specimens have some portion
of the dumbbell-shaped figure present and lack the marginal spots
or have only faint ones. When the dumbbell is reduced, the dis-
appearance of the figure comes at the junction of the pectoral and
abdominal laminae, thus dividing the figure into an anterior and
a posterior spot. The dumbbell may be solid or may have a hollow
center. One specimen (TU 6038) has but a single dot on the bridge,
on the under side of marginal 5.

The markings of the soft parts are typical of the species. Five
lines enter the posterior border of the eye, of which the most dorsal
and most ventral (orbital) are the widest and most prominent, the
central is next in width, and the two intermediate stripes are often
only faintly indicated. In all but one specimen the median head
stripe does not join the canthal line. The ventral markings of the
neck and chin consist of a small median yellow ovoid spot on the
lower beak; in some specimens it is confluent with the median yellow
stripe on the chin and neck. The two paramedian yellow ventral
neck lines begin immediately posterior to the beak in some individ-
uals, and in others they are confluent with the yellow line on the
upper beak. In some specimens these paramedian ventral neck
stripes are confluent with the median triangle on the lower beak.
Thus the neck markings show a variety of conditions, unrelated to
the geographic source of the specimen.

The six smallest individuals (carapace lengths between 29.7 and
35.8 mm.) have the primary or hatchling laminae rugose. The next
group (carapace lengths between 36.5 and 44.5 mm.) has the laminae

474 FIELDIANA: ZOOLOGY, VOLUME 34

smooth or finely striated, while the largest juvenile (carapace length
55.3 mm.) has the carapace completely smooth, with the striae only
faintly indicated. The transition between the rugose and smooth
conditions of the carapace occurs at a carapace length of about
50 mm.

SUBADULTS

Nine subadult individuals show the variation in pattern and
configuration of D. r. reticularia. The carapace lengths vary between
68.3 and 146 mm. The shell outline is distinctly ovoid and not
cuneiform. The ground color of the carapace is very dark (almost
black) in most specimens and the dorsal reticulum and mid-dorsal
yellow line are either very fine or obscured. The ratio of CW to CL
varies between 0.67 and 0.81, the larger individuals having the
smaller ratios. The shell is high in the dorsal median line and may
even be slightly keeled, although this is not the common condition.
The marginal edging is narrow and inconspicuous. The ratio of
D to CW ranges between 0.51 and 0.63.

The plastron is immaculate in six specimens. The remaining
three individuals have some degree of pigmentation on the median
sutural line. One (ChM 43.56) has faint darkening at the juncture
of the femoral and anal laminae; another (ChM 53.115.4) has a wide
(4 mm.) irregular dusky band, most pronounced at the femoral-anal
juncture, extending anteriorly as far as the pectoral laminae. A
third individual (AMNH 46776) has a dusky band 11 mm. wide
extending over the median sutures of the posterior three pairs of
laminae and extending narrowly laterally at the seam of the abdom-
inal and femoral laminae.

In no specimen is the bridge region immaculate. The dumbbell
figure is absent in one individual but expressed variously in the
others, and the spots on the marginals are present in all but one.
There is a tendency for the dumbbell-shaped figure to be joined to
the marginal spots by extensions of the black pigment from the figure
to the spots; the anteriormost spot on marginal 4 is the first spot
to show this joining, and the spot on marginal 5 is the next to be
so joined.

Two specimens have the median head line joined to the canthal
line and in six these two lines do not fuse. The head and neck
striping does not differ from that of the juveniles.

ADULTS

Males. In eleven males the shell is flattened and not deep, and
the outline is roughly ovoid or slightly cuneiform. The dorsal

SCHWARTZ: CHICKEN TURTLE 475

reticulation and mid-dorsal line are usually fine and hardly visible,
but in some specimens they are quite as broad as those of the
Florida subspecies. However, in preserved specimens, the reticu-
lation is gray and not yellow as in the more southern subspecies;
the coloration of these broadly marked turtles in life can only be
conjectured, but since specimens of D. r. reticularia customarily
have a greenish or brownish reticulum rather than an orange or
yellow one as in the Florida turtle, it may be assumed that in life
these aberrant individuals were not so brightly colored. The
ground color of the carapace is dark brown to black, and the yellow
edging of the marginals is very fine or absent. The entire lot displays
a reduction of the carapace markings so that extreme individuals
have these markings much reduced.

Two adult males show a vestige of a plastral blotch at the femoral-
anal suture, but in these two individuals the spot is faint and ill-
defined. One specimen lacks any bridge figure or marginal spots
at the bridge level, and another lacks the marginal spots but has the
dumbbell-shaped figure. The remainder show a variety of con-
ditions; if both the bridge figure and marginal spots are present,
there is a tendency toward fusion of the entire blotch system on the
bridge, thereby forming a dumbbell-shaped figure with blunt dorsad
extensions on the marginals. In some specimens there is fusion
between the marginal spots as well as between the spots and the
dumbbell-shaped figure, resulting in a complex and poorly defined
bridge blotch.

Seven specimens have the median head stripe separate from the
canthal line, while one has these two joined. In the other specimens
the condition is indeterminate.

Females. The twelve females show the extreme reduction of the
dorsal reticulum and mid-dorsal line. The carapace is broadly oval
and high and is colored dark brown to black. The pattern is repre-
sented by only the very finest lines, or hardly at all. The yellow
edging of the marginals likewise disappears.

In two specimens there are faint traces of mid-ventral pigmenta-
tion at the femoral-anal suture, extending sporadically anteriorly
over the abdominal laminae. The remaining specimens have im-
maculate plastra. The bridge lacks figures of any sort on one
specimen, and the dumbbell-shaped figure is barely suggested in
another. Two turtles lack marginal spots at the level of the bridge,
but have the dumbbell figure present and bold. The remaining
specimens have both the figure and the spots variously expressed.

476 FIELDIANA: ZOOLOGY, VOLUME 34

The tendency to fusion of the figure with the marginal spots noted
in adult males is shown here also; in two specimens the figure and
the spots are fused.

The median head stripe is joined to the canthal line in six speci-
mens, separate in three, and indeterminate in three.

Deirochelys reticularia chrysea subsp. nov.

Deirochelys reticularia Loennberg, 1894, Proc. U. S. Nat. Mus., 17: 318;
Stejneger and Harbour, 1917, Check List of N. Amer. Amphibians and
Reptiles, p. 121 (part); Conant, 1930, Bull. Antivenin Inst. Amer., 4
(3), pp. 61, 63; van Hyning, 1933, Copeia, 1: 7; de Sola, 1935, Copeia, 1:
45; Brimley, 1939-43, Reptiles and Amphibians N. Car., Carolina Tips,
Installment 31, second unnumbered page (part); Carr, 1940, Univ. Fla.
Publ., Biol. Sci. Ser., 3 (1), p. 105 (part); Coin, 1943, Proc. Fla. Acad.
Sci., 6: 150; Ditmars, 1946, Reptiles N. Amer., pp. 418-419 (part); Brown,
1950, Check List of Reptiles and Amphibians, Texas, p. 234 (part); Carr,
1952, Handbook of Turtles, pp. 316-319 (part); Telford, 1952, Quart.
Jour. Fla. Acad. Sci., 15 (3), pp. 184-185; Schmidt, 1953, Check List of
N. Amer. Amphibians and Reptiles, p. 104 (part); Carr and Goin, 1955,
Reptiles, Amphibians and Fresh-water Fishes of Fla., p. 246 (part);
Mertens and Wermuth, 1955, Zool. Jahrb. (Syst.), 83, (5), pp. 348-349
(part).

Deirochelys reticulata Siebenrock, 1909, Zool. Jahrb., suppl. vol. 10 (3), p. 485
(part).

Chrysemys reticulata True, 1883, Bull. U. S. Nat. Mus., 24: 35 (part); True,
1884, Fish. Indust. U. S., sec. 1, p. 157 (part); Boulenger, 1889, Cat.
Chel. Brit. Mus., p. 75 (part).

Dierochelys reticularia Loding, 1922, Alabama Mus. Nat. Hist., mus. paper
no. 5, pp. 45-46 (part); Haltom, 1931, Geol. Surv. Ala., Alabama Mus.
Nat. Hist., mus. paper no. 11, p. 137 (part).

Holotype. UMMZ 111440, adult male, taken 5.8 miles east of
Monroe Station, Collier County, Florida, on September 4, 1954,
by L. Neil Bell.

Allotype UMMZ 100371, adult female, taken 5 miles west of
Florida City, Bade County, Florida (no date), by Albert Schwartz.

Paratypes. FLORIDA: St. Lucie Co., 23 miles southwest of Ft.
Pierce (UMMZ 96513); Okeechobee Co., Taylor Creek Slough,
3.5 miles north of Okeechobee (ChM 54.202) ; Highlands Co., Red
Hill, near Hicoria (AMNH 65620), Archbold Biological Station,
near Hicoria (AMNH 65621); Glades Co., 12.4 miles southwest of
Okeechobee (UMMZ 106309); Hendry Co., southeast of La Belle
(UMMZ 106319-20); Collier Co., Immokalee (CAS 9158), 2 miles
west of Immokalee (UF 7219), 7.7 miles south of Immokalee (UF
7220), 2 miles north of Copeland (ChM 55.39.2); Broward Co., New

SCHWARTZ: CHICKEN TURTLE 477

River (MCZ 12497); Dade Co., Tamiami Trail (AMNH 49954),
Miami (AMNH 9054), 5 miles west of Florida City (UMMZ 100370).

The following specimens of D. r. chrysea have been examined
and are not designated as para types: FLORIDA: no other locality, 2;
county indeterminate (not mapped), Colohatchee River, 1; Alachua
Co., Gainesville, 4, 1 mile south of Gainesville, 1, east side of Lake
Newnan, 1, Payne's Prairie, 1, Kale's Siding, 1, Orange Heights, 1,
Monteocha, 3, 5 miles north of Micanopy, 1, no other locality, 3;
Putnam Co., near Welaka, 1, Palatka, 1; Marion Co., 8 miles east
of Ocala, 1, Silver Springs, 2, Dunnellon, 3, 15 miles east of Dun-
nellon, 1, near Levy County line, 1, near Rainbow Springs, 1,
Eureka, 1; Lake Co., Lake Yale, 2, Leesburg, 4, near Leesburg, 1,
1.4 miles north of Leesburg, 1, Eustis, 1; Sumter Co., 2.8 miles
southwest of Wild wood, 1; Orange Co., Winter Park, 1, Orlando, 2;
Brevard Co., 3 miles north of Merritt Island P. O., 1, Eau Gallic, 1,
Titusville, 1, Canaveral, 3; Osceola Co., 2 miles west of Kissimmee,
1, Kissimmee Prairie(?), 1, no other locality, 2; Polk Co., 3 miles
south of Lake Wales, 1, Lowery Lake, 1; Hillsborough Co., Hills-
borough River State Park, 6 miles south of Zephyrhills, 1, 16.6 miles
south of Riverview, 1, west of Plant City, 1, Ruskin, 1, no other
locality, 1; Pinellas Co., Gulfport, 5; Indian River Co., Fellsmere, 1;
Manatee Co., Palmetto, 2; Sarasota Co., Sarasota, 1, Englewood, 1;
Dade Co., Long Pine Key, 1.

Geographic distribution. The Florida peninsula from Alachua,
Putnam, and western Marion counties south to the tip of the
peninsula, exclusive of the Florida Keys. Intergrades between
D. r. chrysea and D. r. reticularia as noted in discussion of the latter
form (see map, fig. 102).

Diagnosis. Deirochelys r. chrysea differs from D. r. reticularia
and D. r. miaria in that the plastron is orange or bright yellow and
lacks a pattern of any sort, the dorsal reticulations are orange or
golden and are bold and broad except in old specimens, and the
orange edging of the carapace is broad and prominent. The black
spots on the ventral surface of the marginals at the level of the
bridge are absent in 57 per cent of the specimens examined.

The shape of the carapace is cuneiform (best noted in males of
all sizes, and juveniles, and least expressed in females), and there is
a constriction at the level of marginals 5 and 6. The carapace of
D. r. chrysea is relatively higher than that of both D. r. reticularia
and D. r. miaria. See Tables 1 and 2.

478

FIELDIANA: ZOOLOGY, VOLUME 34

FIG. 105. Dorsal and ventral views of the holotype of Deirochelys r. chrysea
(UMMZ 111440) from 5.8 miles east of Monroe Station, Collier County, Florida-

Description of holotype. An adult male (fig. 105) with the
following measurements: CL, 110.2; PL, 99.2; C3/4, 63.8; C5/6,
72.9; C8/9, 72.6; CW, 78.6; head width, 20.1; D, 45.8; width of
posterior lobe of plastron, 45.8; width of bridge, 31.9. Ratios:
D/CW, 0.58; C5/6/CL, 0.66; CW/CL, 0.71. The shell is cuneiform
in outline when viewed dorsally and moderately high in the mid-
dorsal line. The greatest width of the carapace is at the level of
marginal 7 and the greatest height at central 3. The marginals
are flaring from marginal 7 posteriorly. There is a marked constric-
tion at the level of marginals 5 and 6 so that the carapace has
a "pinched-in" look at this region. This constricted area plus the

SCHWARTZ: CHICKEN TURTLE 479

flare of the more posterior marginals combine to give the overall
wedge-shaped appearance to the shell.

In the following color description color notes from life have been
utilized whenever possible. Capitalized color names are from
Ridgway (1912). The carapace is Olivaceous Black (1) in color
with the reticulations Tawny Olive. The mid-dorsal line and the
reticulum are prominent and wide (as much as 2.5 mm.). The
carapace is edged with Yellow Ocher, which grades to Buckthorn
Brown posteriorly; this edging is especially prominent at the points
of contact on the marginals with the lateral ends of the reticulations.
The overall effect produced on the marginals is that they are brown-
ish tinged with orange with isolated "islands" of black pigment
at the marginal seams, cut off from the dorsal ground color. The
plastron is immaculate Capucine Yellow. The bridge pattern is
almost absent; there is a small black dot on the left inguinal lamina
and a black blotch on the right. No marginals have ventral spots.
The bridge is Orange on the axillary, inguinal, and marginal laminae
as well as on the lateral extensions of the pectoral and abdominal
laminae.

The head skin is black. Three lines (Aniline Yellow) enter the
posterior margin of the eye; these grade posteriorly on the neck
into Primuline Yellow and Old Gold. The most dorsal and most
ventral (orbital) of these three lines are the widest and most promi-
nent. Two faint yellowish lines enclosed by these two broad lines
represent the usual fourth and fifth lines that enter the margin
of the eye. The median head line is not connected to the canthal
lines, but is connected to one of the five lines on the upper eyelid
and extends posteriorly for about 8 mm. behind the posterior edge
of the eye. The paramedian head lines begin on the upper eyelids
and extend posteriorly to varying distances along the neck. A
post-ocular stripe begins at the posterior corner of the eyelids and
extends as far posteriorly as the margin of the carapace. These
major head lines are usually separated by one faint yellowish line.
The chin has an inverted U-shaped Lemon Yellow blotch, attached
to a black-bordered median bar on the lower beak. The ends of
this figure continue as the major broad yellow bands on the neck.
Between them a third major yellow band begins about 15 mm.
posterior to the rear edge of the lower beak and continues to the
edge of the plastron. There is a Lemon Yellow line on the upper
beak, beginning at the nares and continuing onto the lower beak
as a black-bordered blotch.

480 FIELDIANA: ZOOLOGY, VOLUME 34

The forelimbs have a wide (about 7 mm.) Lemon Yellow band
on the anterior face at the level of digits 2, 3, and 4, and the dorsal
edge of the limb has a yellow line. There is a broad (2 mm.) band
on the ventral edge of the forelimb. The webs are yellow. Between
the elbow and the carapace there are 8 yellow lines on the black
skin. The hind limbs are black dorsally, with the exception of
a yellow band on the dorsal aspect of the thigh. Ventrally, the hind
limbs are yellow (except for the black plantar surfaces) and variously
mottled with black. Between the knee and the vent there are 7
vertical yellow lines on the black skin. The tail is stout, cylindrical,
and tapering (22 mm. from vent to tip), and the vent lies approxi-
mately 16 mm. behind the posterior edge of the plastron. The tail
is yellow laterally and black dorsally, with a yellow mid-dorsal line
on the distal half. Two faint paramedian yellowish lines are in-
dicated on the dorsum of the tail. The penis is extruded and is
black.

Description of allotype. An adult female (fig. 106) with the
following measurements: CL, 187; PL, 169; C3/4, 103; C5/6, 122;
C8/9, 117; CW, 126; head width, 28.0; D, 82; width of posterior
lobe of plastron, 72.3; width of bridge, 55.8. Ratios: D/CW, 0.65;
C5/6/CL, 0.65; CW/CL, 0.67. The shell is oval in outline when
viewed dorsally and is highly domed; the greatest width of the
carapace is at the level of marginal 7 and the greatest height at
central 3. There is a slight mid-dorsal keel. The shell tapers
rapidly anteriorly and is stout and appears to bulge at the level
of lateral 3. The marginals are moderately flaring from marginal 7
posteriorly and marginals 4, 5, and 6 are almost vertical. The
constricted appearance of the holotype and of most males is not
demonstrable in the females of such large size. The carapace is
brown in color and the reticulations and mid-dorsal line are broad
(as wide as 2.5 mm.) and prominent. The carapace is widely
edged with dull orange or yellow, and the juncture of this edging
with the reticulum produces the isolated islands of brown ground
color on the marginal seams as in the holotype. The plastron is
immaculate and orange. The bridge has the usual dumbbell-shaped
figure entire, on the lateral extensions of the pectoral, abdominal,
axillary, and inguinal laminae on the left side, and separated at the
pectoral-abdominal suture on the right. The ventral surface of
the marginals lacks any black blotches or markings.

The head skin is brownish black. Five lines enter the posterior
margin of the eye; of these the dorsal and ventral (orbital) are most
prominent, the median are somewhat less so, and the remaining two

SCHWARTZ: CHICKEN TURTLE

481

FIG. 106. Dorsal and ventral views of the allotype of Deirochelys r. chrysea
(UMMZ 100371) from 5 miles west of Florida City, Dade County, Florida.

are fainter still. The median line is not joined to the canthal line,
and extends far back on the neck (which is hardened in position and
cannot be extracted from the shell). The paramedian lines begin
about 5 mm. behind the eye. The details of head striping are much
as in the holotype, except that usually 3 faint lines separate the
major head stripes. The throat pattern is asymmetrical and consists
of two lateral yellow bands which begin just posterior to the lower
beak. The left band turns dorsally and ends at the angle of the jaws;
the right ends posteriorly to form the right major band on the neck.
The left major neck band begins posterior to the angle of the jaws
and continues posteriorly with its mate. There is no well-defined
median ventral neck band. The coloration and pattern are es-
sentially those of the holotype; there are 9 vertical bands between
the knee and the vent, however. The tail is long and terete (32
mm. from vent to tip) and the vent is located 14 mm. behind the

482 FIELDIANA: ZOOLOGY, VOLUME 34

posterior margin of the plastron. The tail is yellow ventrally and
black dorsally, with a pair of paramedian yellow stripes on the
dorsum; the stripes join to form a single median dorsal yellow line
on the distal half of the tail.

JUVENILES

Fifteen juveniles of Deirochelys r. chrysea demonstrated the
characters of this subspecies. The carapace lengths of this series
range between 31.6 mm. and 79.0 mm. The ratio of CW to CL
varies between 0.81 and 0.96, with generally the lower ratios for the
larger specimens. The shell shape, other than in the very smallest
individuals, is markedly cuneiform, with the posterior half of the
carapace widely flaring. The carapace is brown dorsally, and the
yellow reticulum and mid-dorsal line are wide and boldly prominent.
The margin of the carapace is widely edged with orange or yellow.
The carapace is high mid-dorsally and the ratio of D to CW varies
between 0.50 and 0.60. The orange edging of the carapace may
involve as much as one- third of the marginal laminae posteriorly,
and when the edging is so broad, it joins the yellow reticulation
broadly on each marginal scute. In one individual (UF 2906) the
dorsal reticulation is faint, but present.

The plastron is immaculate and yellow (orange in fresh material)
in all specimens. The bridge pattern varies between a complete
absence of any markings on either marginals or the bridge laminae
proper (USNM 131832, ChM 54.237.4) to a completely expressed
pattern of the usual dumbbell-shaped figure on the inguinal and
axillary laminae and on the lateral extensions of the pectoral and
abdominal laminae. Of the thirteen specimens with some markings
on the bridge region, seven lack any marginal black dots, and an
eighth has these only feebly expressed. The dumbbell-shaped
figure may be reduced to merely a small blotch on the inguinal
lamina. There is thus a tendency toward reduction of the bridge
markings, and this tendency is clearly shown in the entire lot of
D. r. chrysea.

The coloration of the soft parts is typical of the species. There
are five lines entering the posterior margin of the eye, and although
these may be of equal width (CNHM 46273) usually the median,
dorsal, and ventral (orbital) stripes are the most prominent. In all
specimens save one (AMNH 65620) the median head stripe does not
join the canthal line.

The chin and neck markings are complex and variable but
typically consist of a median mental blotch on the lower beak,

SCHWARTZ: CHICKEN TURTLE 483

surrounded by a triangular figure which is produced posteriorly
into a median yellow line on the chin. Two broad yellow stripes
begin at the postero-lateral sides of this triangle and continue
posteriorly as the conspicuous broad yellow lines on the neck. In
some specimens these two broad lines are joined laterally and dor-
sally to the yellow or orange line on the upper beak.

In the three smallest specimens (carapace lengths 31.6 to 36.3
mm.) the carapace laminae show the rugose condition of the primary
laminae. In specimens of carapace lengths between 45.6 and 69.4
mm. new growth is smooth, and the rugose primary laminae are
clearly distinct from the secondary laminae. Specimens of carapace
lengths between 59.0 and 79.0 mm. have lost the shagreened ap-
pearance of the carapace laminae and these plates are now either
smooth or show the fine striae typical of the adult. The transition
between these two conditions occurs in D. r. chrysea in turtles
between 60 and 70 mm. in carapace length.

SUBADULTS

Eighteen subadults (carapace lengths between 79.3 and 141.1
mm.) show the amplification of the trends demonstrated in the
juveniles. The carapace continues to develop in a wedge shape and
marginals 5 and 6 are almost vertical while marginals 7 through 9
are widely flaring. The ratio of CW to CL varies between 0.68 and
0.80, while the ratio of D to CW varies between 0.44 and 0.62. The
reticulum is broad and prominent on a dark brown background, and
the mid-dorsal line may be either prominent or obscured to a fine
line. The yellow edging of the carapace is prominent and may be
quite broad. Compared to D. r. reticularia at the same stage of
development, the dorsal pattern is far more prominent and the
reticulum is wider in the present form.

The plastron is immaculate in all specimens. The bridge lacks
the dumbbell-shaped figure as well as the marginal markings in one
specimen, and eleven specimens have only the figure and lack the
marginal blotches at the bridge. The remaining six specimens have
some indication of the dumbbell-shaped figure and the marginal
blotches, although both may be merely suggested. The marginal
blotches are usually, when fully expressed, complete circles (some-
times with hollow centers), restricted either to a single marginal
lamina or encroaching slightly upon the next posterior lamina.
The dumbbell figure may be solid or practically hollow. When the
bridge figure is reduced, it may remain as two dark smudges over-

484 FIELDIANA: ZOOLOGY, VOLUME 34

lying the axillary-pectoral and inguinal-abdominal sutures, re-
spectively.

The pattern of the head, chin, and neck is as described for the
juveniles. One individual has the median head line joined to the
canthal line.

ADULTS

Males. Twenty-eight males are available, and this series shows
the variation in adult pattern and size in male D. r. chrysea. In
general the cuneiform outline of the shell is preserved except in
old males, which are more ovoid in carapace shape. The dorsal
reticulations and mid-dorsal line are usually prominent, but in a few
individuals they are less so. There seem to be two general variants
in ground color of the carapace. One group consists of individuals
in which the ground color is light brown to olive in color, and the
reticulum is broad and persistent. In other individuals the ground
color is dark brown or almost black, and here the reticulum is often
somewhat obscured, due to darkening of the orange pigments of
the reticulum as well as encroachment of the carapace pigment on
to the reticulum. However, in most cases the reticulations and the
mid-dorsal line are quite obvious.

One specimen has the bridge completely immaculate, and
fifteen lack the marginal spots on the bridge. In the remainder
the dumbbell-shaped figure is variously expressed, and the largest
individual (UF 534) shows obsolescent hollow smudges on the bridge
at the axillary-pectoral and inguinal-abdominal sutures. This may
indicate that at very large sizes the bridge markings of males become
obsolete.

The head markings are as already noted; three individuals have
the median head stripe joined to the canthal line, while the remainder
of the specimens have this median line separate and unjoined.

Females. Except in very large and old individuals the twenty
adult females show that the reticulations and the mid-dorsal line
remain distinct and brightly colored. The edging of the carapace
is prominent and very wide; the allotype represents an extreme in
this regard since the yellow edge to the marginals extends broadly
upon the marginals, and the juncture of this edge with the termi-
nations of the reticulum upon each marginal results in cutting off
the almost circular blotches of the dorsal ground color at the inter-
marginal sutures.

In no specimen does the median head line join the canthal line.

SCHWARTZ: CHICKEN TURTLE 485

The carapace is as usual immaculate. Two individuals have the
bridge completely immaculate, while the marginal spots are lacking
in eleven. The remainder have the bridge pattern variously ex-
pressed.

Intergradation between D. r. reticularia and D. r. chrysea occurs
in a relatively narrow band across the Florida peninsula just north of
Alachua County in the central and eastern portion of the state, and
in Levy County on the western seaboard. Intergrades have been
examined from the following Florida localities: Taylor Co., 17 miles
southeast of Perry, 1; Dixie Co., 1.5 miles southeast of Cross City, 1,
3.2 miles southeast of Clara, 1; Levy Co., 10 miles south of Chief-
land, 1, 7 miles northwest of Otter Creek, 1, 4.7 miles northwest of
Otter Creek, 1; Gilchrist Co., no other locality, 1; Clay Co., no other
locality, 1. These few specimens are intermediate between D. r.
reticularia and D. r. chrysea in that the color of the carapace begins
to approach the black or dark brown coloration of D. r. reticularia
and the dorsal reticulum is progressively more narrow and incon-
spicuous in specimens collected from south to north. In this latter
character, the specimens from Dixie and Levy counties are closer to
D. r. chrysea (the reticulum is broad), but the individuals from
Taylor, Clay, and Gilchrist counties are closer to D. r. reticularia.
The plastral color, which is also diagnostic between these two
subspecies, cannot be determined from this preserved material, and
no living examples from the area have been studied.

Occasional specimens from the geographic range of D. r. reticu-
laria resemble D. r. chrysea in shape of carapace and width of
reticulum. A single individual from southern Mississippi and two
from west-central South Carolina deserve mention in this regard.
These specimens have been designated herein as D. r. reticularia
in spite of this resemblance to D. r. chrysea and serve to emphasize
the close relationship between the two subspecies occurring east
of the Mississippi River.

In addition to the holotype of D. r. chrysea, color notes have
been taken from two live specimens of this subspecies (adult male
from 3.5 miles north of Okeechobee, Okeechobee Co., Fla.; juvenile
from Lake Yale, Lake Co., Fla.). Color notes from eight live speci-
mens of D. r. reticularia have also been obtained (adult female from
Lake Moultrie near Pinopolis, Berkeley Co., S.C.; subadult female
from Santee National Wildlife Refuge, Lake Marion, Clarendon
Co., S.C.; three adult females from 2.1 miles north, thence 1.2 miles
west of Cainhoy, Berkeley Co., S.C.; juvenile from 0.9 miles north

486 FIELDIANA: ZOOLOGY, VOLUME 34

of Cainhoy, Berkeley Co., S.C.; adult female from 11 miles west
of Bogalusa, Washington Parish, La.; adult male from 2.1 miles
north, thence 1.2 miles west of Cainhoy, Berkeley Co., S.C.). Colors
of these ten specimens are presented in Tables 2 and 3; colors are
tabulated for each specimen in the order as listed above.

Inspection of Tables 3 and 4 shows that the colors of the Flori-
dian subspecies are in general brighter than those of D. r. reticularia.
This is especially true of the color of the plastron and the bridge.
The skin markings are also somewhat paler in D. r. reticularia, and
in this race the dorsal head and neck markings show a pronounced
tendency toward greenish in contrast to the bright yellows of D. r.
chrysea. Unfortunately, adequate series of live D. r. chrysea are not
available for study and comparison with D. r. reticularia, and no
attempt has been made to assess differences in coloration between
the sexes (males appear to be somewhat more brightly colored than
females) and between juveniles and adults, because of the meager
material available. The data here presented indicate that there are
differences in coloration between the peninsular and mainland
populations of Deirochelys.

Wright and Funkhouser (1915, pp. 117-118) described the plas-
tral coloration of an adult chicken turtle from the Okefinokee
Swamp as "reddish-yellow, median plates with a distinct tinge of
red" and that of a juvenile as "clear lemon yellow." It is possible
that the brightly colored plastron typical of D. r. chrysea occurs in
individuals as far north as the Okefinokee Swamp region of southern
Georgia, but on the basis of shell outline and proportions, material
from the Okefinokee Swamp is assigned to subspecies reticularia.
Additional fresh material from this area may show that south-
eastern Georgia is an area of intergradation between D. r. reticularia
and D. r. chrysea.

Deirochelys reticularia miaria subsp. nov.

Deirochelys reticularia Bellinger and Black, 1938, Occ. Pap. Univ. Arkansas
Mus., no. 1, p. 45; Schwardt, 1938, Univ. Arkansas Coll. Agric., Bull. 357,
p. 45; Pope, 1939, Turtles U. S. and Canada, pp. 234-235 (part); Brimley,
1939-43, Reptiles and Amphibians N. Car., Carolina Tips, Installment 31,
second unnumbered page (part); Ditmars, 1946, Reptiles N. Amer.,
pp. 418-419 (part); Brown, 1950, Check List of Reptiles and Amphibians,
Texas, p. 234 (part); Webb, 1950, Herpet., 6 (5), pp. 137-138; Carr, 1952,
Handbook of Turtles, pp. 316-319 (part); Smith and Sanders, 1952, Texas
Jour. Sci., 4: 211; Guidry, 1953, Herpet., 9 (1), p. 56; Schmidt, 1953,
Check List of N. Amer. Amphibians and Reptiles, p. 104 (part); Carr and
Goin, 1955, Reptiles, Amphibians and Fresh-water Fishes of Fla., p. 246

SCHWARTZ: CHICKEN TURTLE 487

(part); Mertens and Wermuth, 1955, Zool. Jahrb. (Syst.), 83 (5), pp.

348-349 (part).
Deirochelys reticulata Agassiz, 1857, Contr. Nat. Hist. U. S., 1: 441 (part),

2, pi. 2, figs. 1-3; Siebenrock, 1909, Zool. Jahrb., suppl. vol. 10 (3), p. 485

(part).
Chrysemys reticulata True, 1883, Bull. U. S. Nat. Mus., 24: 35 (part); True,

Fish. Indust. U. S., sec. 1, p. 157 (part).

Holotype. CNHM 37478, adult male, taken at College Station,
Brazos County, Texas, on April 17, 1941, by K. P. Schmidt, J. M.
Schmidt, C. M. Barber, and A. Flury.

Allotype. USNM 85145, subadult female, taken at White Rock
Creek, 4 miles northeast of Dallas, Dallas County, Texas, on April
16, 1932, by C. E. Burt.

Paratypes. TEXAS: Dallas Co., 6 miles southeast, Y^ mile east
of Trinity River (TNHC 8907); Freestone Co., 20 miles east of
Teague (BCBC 4694) ; McLennan Co., south of Waco (BCBC 4690),
5 miles east of Waco (SM 157), 3 miles east of Axtell (SM 0085,
160), Trading Horse Pond (SM 6684), 1 mile southeast of Harrison
Switch, Kirkpatrick Lake (CNHM 52995); Leon Co., 5 miles west
of Marquez (CNHM 46282-3, 52996) ; Brazos Co., College Station
(TCWM 4680), Green Lake (TCWM 691); Grimes Co., 7 miles
northeast of Navasota (TCWM 7278) ; Liberty Co., 3.7 miles north
of Hull (TNHC 8906); Austin Co., 11 miles northwest of Sealy
(BCBC 225), San Felipe (CNHM 37482); Harris Co., 1.2 miles
southeast of Bammel (TNHC 6035), Houston (CM 5443-5), South
Houston (ChM 55.39.3, TNHC 8905) ; Colorado Co., 8 miles from
Eagle Lake (TCWM 326), Rock Island (UK 3145); Ft. Bend Co.,
3 miles south of Beasley (BCBC 3044-6, 3048, TCWM 4679,
UIMNH 2397), Fresno (UIMNH 28503, 31696), 1 mile west of
Orchard (AMNH 36726) ; Lavaca Co., 22 miles southeast of Yoakum
(TNHC 7325).

The following specimens of D. r. miaria have been examined
and are not designated as paratypes: ARKANSAS: Faulkner Co., near
Con way, 1; Craighead Co., 6 miles west of Jonesboro, 1; Jackson
Co., 3 miles south of Swifton, 1; Prairie Co., Duvall's Bluff, 1;
Lonoke Co., 2.6 miles east of Carlisle, 1, Lonoke, 1; Drew Co.,
Jerome, 2. OKLAHOMA: Cleveland Co., 15 miles east of Norman, 1;
Seminole Co., Bowlegs, 1; McCurtain Co., no other locality, 1.
TEXAS: Hopkins Co., 2 miles west of Saltillo, 2; Harrison Co.,
5 miles northeast of Karnack at Caddo Lake, 1; Chambers Co., 5
miles east of Anahuac, 1 (not mapped); Jefferson Co., 2 miles
west of Sabine Pass, 1. LOUISIANA: no other locality, 1; Morehouse

488 FIELDIANA: ZOOLOGY, VOLUME 34

Parish, Mer Rouge, 1, Prairie Mer Rouge, 2 (not mapped) ; Richland
Parish, Rayville, 5; Franklin Parish, Wisner, 3; Concordia Parish,
Ferriday, 6; Jefferson Davis Parish, 2 miles west of Welsh, 1;
Acadia Parish, Rayne, 9; Orleans Parish, New Orleans, 6; parishes
indeterminate (not mapped), Urosius, 1, Bayou, 2, south central
Mississippi Valley, 6.

Geographic distribution. West of the Mississippi River in
Louisiana, southern and eastern Arkansas, southeastern Oklahoma,
and eastern Texas (see map, fig. 102).

Diagnosis. Deirochelys r. miaria differs from both D. r. reticu-
laria and D. r. chrysea in possessing a plastral pattern of the seam-
following type at all ages; this pattern is usually present but
obsolescent in old adults. Adults usually have an unstreaked chin
and throat.

The carapace is oval in shape and flatter than that of both
D. r. reticularia and D. r. chrysea. See Tables 1 and 2.

Description of holotype. An adult male (fig. 107) with the
following measurements: CL, 144; PL, 130; C3/4, 87; C5/6, 104;
C8/9, 100; CW, 111; head width, 22.8; D, 56; width of posterior
lobe of plastron, 62.3; width of bridge, 42.0. Ratios: D/CW, 0.50;
C5/6/CL, 0.72; CW/CL, 0.77. The shell is oval in outline when
viewed dorsally and is quite flattened. The greatest width of the
carapace is at the level of marginal 8 and the greatest height at
central 2. The posterior marginals flare moderately from marginals
7 to 9, but this is not so noticeable in the posteriormost marginals.
There is no constriction at the level of marginals 5 and 6 as in
D. r. chrysea, and these marginals slope medially, not vertically.
The reticulum and mid-dorsal line are wide (almost 3 mm. broad)
and, although obvious, the color is only slightly lighter than the
olivaceous ground color of the carapace. The carapace is incon-
spicuously edged with yellow on the sides, and no yellow border
occurs on the anterior or posterior marginals. The terminations
of the reticulum continue to the edge of the carapace and are slightly
widened at the very edge of the shell. The plastron is yellow, with
heavy deposition of dark pigments. The age of this individual
accounts for the deposition of brown pigments over the entire
surface of the anal laminae, and the central portions of the humeral
and abdominal laminae. Anteriorly, brown pigments follow the
median and lateral seams, and the gular, humeral, and pectoral
laminae are diffusely mottled with brown. The bridge pattern
consists of the entire dumbbell-shaped figure as well as dark brown

SCHWARTZ: CHICKEN TURTLE

489

FIG. 107. Dorsal and ventral views of the holotype of Deirochelys r. miaria
(CNHM 37478) from College Station, Brazos County, Texas.

blotches at the seams between marginals 3-7, on the left side, and
between marginals 2 and 3 and 7 and 8, on the right. The ventral
surface of the marginals from marginal 8 around the posterior
circumference of the shell is clouded with brown.

The head skin is dull tan (faded?). Two major yellowish lines
enter the posterior margin of the eye; between them are faint
indications of the usual three enclosed lines, the median of which
is slightly more prominent. The median line is obscured but appears
not to have been joined to the canthal line. Two paramedian lines
begin about 3 mm. posterior to the eye and extend posteriorly along
the neck to the carapace. The post-ocular stripes begin about 5 mm.
behind the eye and continue as major neck stripes to the carapace
edge. There is no indication of striping between these major head
and neck lines. The chin and ventral surfaces of the neck are

490 FIELDIANA: ZOOLOGY, VOLUME 34

practically unstreaked. The usual ventral neck markings begin
at about the level of the tympanum and continue inconspicuously
to the margin of the plastron. The lower beak is flecked with black
along its edge and is elsewhere clear yellow, while the chin and
anterior ventral surface of the neck are yellow, with very faint
indications of the throat streaking typical of the species.

The forelimbs have a broad (12 mm.) yellow band covering
two-thirds of the anterior face and the lower edge of the limb; the
upper edge likewise has a yellow line. Digits 3 to 5 are yellow, as
are the webs also. Between the elbow and the carapace there are

7 vertical yellow lines on the tan ground color. The hind limbs
are tan dorsally and yellow ventrally, except for the plantar surfaces,
which are tan. A yellow stripe extends along the medial aspect
of the limb and foot. Between the knee and the vent there are 7
vertical yellow lines on the tan skin. The tail is stout, cylindrical,
and tapering (41 mm. from vent to tip) and the vent lies about
17 mm. posterior to the posterior edge of the plastron. The tail
is yellow ventrally and brown dorsally, with a median yellow line
on its distal half.

Description of allotype. A subadult female (fig. 108) with the
following measurements: CL, 134.6; PL, 122.7; C3/4, 81.5; C5/6,
94.9; C8/9, 92.5; CW, 100.4; head width, 21.7; D, 50; width of
posterior lobe of plastron, 60.0; width of bridge, 42.7. Ratios:
D/CW, 0.50; C5/6/CL, 0.71; CW/CL, 0.75. The shell is oval and
flattened in outline when viewed dorsally. The greatest width of
the carapace is at the level of marginal 7 and the greatest height
at central 3. The posterior marginals flare moderately from mar-
ginal 8 and the lateral marginals are almost vertical. The dorsal
ground color is tan and the reticulations are very fine and almost
invisible. The carapace is inconspicuously edged with yellow.
The carapace laminae are somewhat translucent and the sutures
of the underlying bones are apparent. The plastron is yellow with
the completely developed seam-following brown plastral pattern.
This pattern involves the entire central seam except for the inter-
gular portion and the extreme posterior quarter of the inter-anal
suture, and all except the lateralmost portions of the remaining
plastral seams. At its widest, the brown pigment presents an area

8 mm. wide on the anal laminae; it is absent on portions of the
inter-abdominal seam. On all lateral seams, the pigment lies on the
posterior margin of the preceding lamina with the exception of
the femoral-anal seam, where two small (4 mm.) circles of pigment
lie at the end of the pigmented seam pattern and project posteriorly

SCHWARTZ: CHICKEN TURTLE

491

FIG. 108. Dorsal and ventral views of the allotype of Deirochelys r. miaria
(USNM 85145) from White Rock Creek, 4 miles northeast of Dallas, Dallas
County, Texas.

onto the anal laminae. The bridge has the complete dumbbell-
shaped figure, which is somewhat hollowed centrally. The marginals
have a brown circular spot at all the marginal seams except at the
junction of the precentral and marginal 1.

The tan head skin shows the same basic pattern as described
for the holotype although it differs in some details. There are five
rather than two major lines entering the posterior margin of the
eye, but the dorsal and ventral (orbital) are the major and most
prominent of these. The median head line is unattached to the
canthal lines. There is but one faint yellowish line between the
major head lines. The post-ocular lines begin at the eye. The throat,
chin, and ventral surface of the neck are immaculate yellow and
show no trace of streaking. The lower beak is yellow and slightly
mottled with brown along its free edge. The pattern and coloration

492 FIELDIANA: ZOOLOGY, VOLUME 34

of the remaining soft parts are essentially the same as those of
the holotype.

JUVENILES

Thirty-eight juveniles of Deirochelys r. miaria are available.
These range in carapace length from 30.8 to 77.8 mm. The very
young individuals (fourteen, with carapace length between 30.8
and 37.1 mm.) are almost circular in outline, the ratio of CW to CL
varying between 0.90 and 0.96. In all the carapace is olive in color-
ation, with the customary yellowish reticulum. There is a moder-
ately well-developed mid-dorsal yellow line. The margin of the
carapace is characterized by a more or less well-developed yellow
to orange border; in one individual (TU 6355, no. 3) the edge of
the carapace is extensively bordered with yellow so that the olive
dorsal coloration along the marginal seams is reduced to dagger-like
points and on marginals 4 to 7 the encroachment of the yellow pig-
ment has caused the formation of olive circular "islands" on the
marginals. No other specimen is quite so extreme.

The plastron is yellow and the black plastral pattern is of the
seam-following type. At its maximum, this pattern involves the
median sutures between the humeral, pectoral, abdominal, femoral,
and anal laminae, and extends laterally on the sutures between the
gular and humeral, humeral and pectoral, pectoral and abdominal,
abdominal and femoral, and femoral and anal junctures. These
lateral branches lie on the posterior margin of each lamina. The
pattern does not reach the bridge at any point. At the union of
the femoral and anal laminae there is an increase in width of the
black pattern, and this customarily extends posteriorly along the
median inter-anal suture. There is a lateral expansion of pigment
at the tips of the branch of pigment along the femoral-anal suture,
extending posteriorly onto the anal lamina. One individual (USNM
17) lacks the plastral pattern, but another specimen from the same
locality has it fully expressed.

On the lateral portions of the pectoral and abdominal laminae
the bridge is marked by a dumbbell-shaped figure that encroaches
onto the inguinal and axillary laminae. This figure is usually
entire, but it may be divided into two or three separate centers of
pigmentation or may be reduced to an anterior and a posterior
blotch; it is not absent on any specimen. The marginals have, at
the level of the bridge, a black spot at each suture; these spots lie on
the marginal immediately anterior to the suture. Two individuals
lack these marginal spots.

SCHWARTZ: CHICKEN TURTLE 493

The soft parts are typical of the species. The markings of the
chin and neck are complex, but their simplest form consists of an
inverted V, with the apex on the beak of the lower jaw and extending
posteriorly along the ventral side of the neck. Three or five lines
enter the eye; if five are present, these are three major lines (most
ventral is the orbital) plus two additional fine lines between the
three major ones.

The carapace laminae are rugose, the rugae on the anterior
portion of the central and lateral laminae being arranged to form
weak striations.

The remainder of the juvenile individuals (carapace lengths
between 35.1 and 77.8 mm.) resemble the younger individuals in
most details of coloration. The plastral pattern is well developed
in all specimens, and there is a tendency for the median portions
of the figure to be broader than the lateral branches. If the lateral
branches of the figure are lost, the branches between the gular and
pectoral laminae disappear first. The marginal spots are generally
restricted to the marginal laminae anterior to the suture, but in
several individuals these spots are expanded onto the marginal
lamina immediately posterior to the suture. Likewise the dumbbell-
shaped figure shows a slight tendency to become hollow, with either
an anterior and posterior light area on the lateral edges of the pec-
toral and abdominal laminae, or a continuous light area within the
entire figure.

Perhaps the most striking change at this period is the disappear-
ance of the markings on the throat and ventral neck. The black
lines are obsolescent on the ventral portion of the neck in an in-
dividual with a carapace length of 54.5 mm. and are completely
lacking in an individual 53.7 mm. in carapace length. The neck
is thus uniformly yellow. Of the lot of twenty-four individuals
under consideration, twelve have either a clear yellow throat or
the black stripes are faded and obsolescent. However, one of the
largest of the lot (carapace length 75.7 mm.) retains the black neck
markings. Although the throat and neck may be clear yellow, the
black markings are retained on the lower beak.

The ratio of CW to CL at this size varies between 0.80 and
0.92, with the smaller individuals having the higher ratios. The
carapace has become more oblong and in most individuals is paral-
lel-sided, with little posterior flare. The aspect of the turtle is
depressed; the reticulations on the carapace may be almost absent
and barely indicated, or they may be prominent. There may be

494 FIELDIANA: ZOOLOGY, VOLUME 34

a faint mid-dorsal line indicated, but it is never prominent as in the
very young individuals.

The texture of the carapace laminae is no longer rugose as in
the very young specimens, but is now smooth or but weakly striated,
foreshadowing the striated condition in the adults. An individual
(CM 24565) with a carapace length of 42.9 mm. shows the smooth
new growth about the primary carapace laminae, in strong contrast
to the original hatching laminae, which are markedly rugose. With
increasing size, the rugosity of the original laminae becomes more
obscure, and it disappears completely in specimens with carapace
lengths exceeding 50 mm.

SUBADULTS

Seventeen individuals (carapace lengths between 79.7 and 139
mm.) are considered to be subadult. The carapace is olive in color
and extremely flattened for Deirochelys; the sides are almost parallel
for the distance between the fourth and seventh marginals. The
ratio of CW to CL varies between 0.72 and 0.81. The dorsal reticu-
lum is almost obsolete and hardly evident. A faint mid-dorsal line
is sometimes present. In some individuals the laminae are almost
translucent, the bone sutures below the centrals being clearly
visible.

The plastral figure is still apparent but some individuals show
the beginnings of the expansion of the figure. The smallest specimen
(UIMNH 31696) to show this melanistic tendency measures 96.8
mm. in carapace length. In this specimen the dark pigment has
expanded laterally from the central seam onto all the plastral
laminae except the gulars and pectorals and is especially prominent
at the junction of the abdominals and humerals. Two other turtles
(carapace lengths of 131 and 139 mm.) also show the same phenome-
non, but the pigment is less extensive, occurring principally on the
posterior half of the plastron. The remainder of the series (of
intermediate size between these incipiently melanistic individuals)
has the customary plastral pattern. There are no concomitant
changes in the coloration of the soft parts or the carapace.

With but two exceptions, the throats and necks of this lot of
subadults are clear yellow. One of these exceptions is the small
individual mentioned above as melanistic, and other (USNM 95345)
is a crushed individual with a plastral length of 106.6 mm.

The spots on the ventral surface of the marginals are either
completely circular or semicircular, the intermarginal sutures limit-
ing the posterior development of the circles.

SCHWARTZ: CHICKEN TURTLE 495

ADULTS

Males. Twenty-three males show the continuation of the
changes indicated in the subadult group of specimens. The cara-
pace retains its flattened appearance, and the sides of the shell
are almost parallel. The dorsal reticulum and the mid-dorsal line
may be almost absent or quite wide, and this condition is not
correlated with the provenance of the specimen or with age. The
melanistic trends noted in subadults are expressed more fully in
adult males; almost all individuals in this group show some modi-
fication of the plastral figure. It may be poorly expressed and
represented by a dark smudge or blotch at the juncture of the anal
and humeral laminae and absent on the anterior portion of the
plastron; or it may be fully expressed on all the plastral laminae,
with increasing pigmentation occurring along the median seams.
The largest male (UOMZ 10940) has a carapace length of 161 mm.
and has the plastral pattern restricted to an irregular blotch on the
posterior three pairs of laminae; the remainder of the plastron is
virtually immaculate and there is no seam-following figure.

Of this lot of males, eleven have a clear yellow or only faintly
streaked throat, while the remainder have a heavily streaked throat.
This is correlated with neither age nor geography.

Females. Sixteen females resemble the males in details of cara-
pace coloration. With increasing size the flatness of the turtle
becomes less obvious, and younger individuals show the more
flattened condition. The old adults have the high-domed shell
which is characteristic of female Deirochelys, and the animal is ovoid
in appearance.

Only one female has the plastral figure expressed completely
and without the additional black pigment. Usually the plastral
pattern is absent over the entire plastron except for an irregular
smudge at the intersection of the humeral and anal laminae, and
even this smudge is lacking in one specimen.

All but one female have clear unstreaked throats.

I have examined one live specimen of D. r. miaria, a subadult
female taken 5 miles east of Anahuac, Chambers County, Texas. The
coloration of this individual in life was as follows: carapace, Dark
Citrine; reticulum, Orange-Citrine and very obscure; carapace edge
Aniline Yellow; plastron Buff- Yellow, pattern black; bridge Apricot
Yellow, grading to Light Cadmium on ventral surface of marginals
2 to 4 and upon axillary lamina. Coloration of beak Martius
Yellow; dorsal head and neck markings Oil Yellow; lower orbital

496 FIELDIANA: ZOOLOGY, VOLUME 34

line and chin Picric Yellow, chin and anterior portion of ventral
neck skin completely unstreaked; tail, hind limb, and rump lines
Picric Yellow; forelimb band Picric Yellow.

Deirochleys r. miaria occurs on the west Gulf coastal plain and
its eastern border is the Mississippi River. It is not surprising that
this large river acts as a boundary between two subspecies of the
aquatic chicken turtle; Carr (1952, p. 318) has pointed out that
Deirochelys "is essentially an inhabitant of quiet water ponds,
marshes, sloughs, and ditches. It is only rarely found in streams,
and then usually in the quieter reaches or arms where pond con-
ditions are approached." However, from the lower reaches of the
Mississippi River there are fourteen anomalous or questionable
specimens which require comment, as follows:

(1) UMMZ 96516, from 20 miles north of Baton Rouge, La. This
individual, if the locality is correct, is from the east side of the
Mississippi River. It is, however, typical of the subspecies miaria in
that it possesses a low, flattened, olive-colored carapace, unmarked
chin and neck, and plastral pattern. Baton Rouge lies on the
Mississippi River and it is conceivable that this specimen is from
west, rather than north of the city, as the locality data indicate.
It is not here interpreted as an occurrence of D. r. miaria east of the
Mississippi River.

(2) UF 874, from near Plaquemines, La. Plaquemines lies on
the west bank of the Mississippi River, and this individual is
a juvenile typical of D. r, reticularia, even possessing the plastral
spot at the juncture of the femoral and anal laminae. Again, the
specimen may well have come from the eastern side of the river and
the locality is here interpreted as incorrect.

(3) CU 1750, from Plaquemines, La. This specimen is typical
of D. r. reticularia and the comments above (see 2) apply equally
well to it.

(4) UF 1737 (2), from 6 miles from Luling, La. Luling lies
on the west bank of the Mississippi River. Since no direction is
indicated from this village, it is conceivable that these two juvenile
individuals are from the east rather than the west side of the river.
They are typical D. r. reticularia and lack any semblance of the
plastral pattern of D. r. miaria.

(5) UIMNH 94-96 and 98-100, from New Orleans, Orleans
Parish, Louisiana. This lot of six juveniles is typical of D. r. miaria;
all have the completely expressed plastral pattern and low carapace

SCHWARTZ: CHICKEN TURTLE 497

of that form. New Orleans is located on the north (the east) bank of
the Mississippi; the present city limits, however, include a portion
of the south (the west) bank of the river. It is presumed that these
six specimens came either from that portion of the city which lies
on the west side of the Mississippi River or from the west bank
of the river "in the vicinity" of New Orleans. These specimens are
designated as D. r. miaria and have been included in the discussion
of variation in that form.

(6) CU 1298 (3), from Schriever, Terrebonne Parish, La. Schrie-
ver is situated about 18 miles south of the Mississippi River on the
west side of that river near Thibodaux. The three specimens are
juveniles. Two have the completely expressed plastral pattern of
D. r. miaria while the remaining individual has at the femoral-anal
suture the plastral spot characteristic of D. r. reticularia. Whether
this situation indicates intergradation between D. r. reticularia and
D. r. miaria cannot be determined on the basis of these three juvenile
specimens; furthermore, intergradation at this locality seems un-
likely on the basis of the other specimens from the Mississippi
delta region. The patternless individual is here interpreted as being
a D. r. miaria that lacks a complete plastral pattern; the remaining
two typical individuals from the same locality tend to bear out
this interpretation.

DISCUSSION

Except for three isolated records, Deirochelys reticularia is
limited in its distribution to the coastal plain. Throughout its
range, the chicken turtle inhabits ponds, lakes, and ditches in
general, lenitic situations and has not been recorded from fluviatile
situations. Of the specimens examined for the present study,
chicken turtles from two rivers (judged by the locality data only)
are at hand. These include specimens from the Tombigbee River
in Alabama and the Chipola River in western Florida; although
the data on these specimens indicate specifically that these turtles
were river-taken, it is of course possible that they were collected
in ponds in the immediate vicinity of the rivers, or from lenitic
backwaters. In general, however, Deirochelys is not a river in-
habitant; in Florida and South Carolina, where I have had experi-
ence with this turtle, all specimens were secured from standing water
(or water with a very low flow gradient) or from fluctuating pools
such as cypress ponds. It is not surprising that the Mississippi
River has proven an effective barrier between the subspecies reticu-
laria and miaria, for this great river appears to be uninhabited by

498 FIELDIANA: ZOOLOGY, VOLUME 34

chicken turtles, and, with the possible exception of its lowermost
reaches, thus acts as a barrier to prevent gene flow between the
populations on the eastern and western banks of the river.

The three extra-coastal plain records of occurrence of the chicken
turtle are from the states of Oklahoma and Arkansas. Webb (1950)
has discussed this occurrence of Deirochelys in Oklahoma; the two
non-coastal plain specimens (Cleveland and Seminole counties) are
assumed to have reached this region via the Mississippi and Arkansas
rivers. Webb stated (1950, p. 137) that "the migration of Deiro-
chelys by way of rivers is incongruous with the habits given by
Pope, . . . but it is possible that the preferable habitat of still water
ponds could present itself" since "the combination of the torrential
rains and the easily eroded sandstone . . . has resulted in the for-
mation of numerous sloughs and ponds, of which the larger are often
relatively permanent." The specimen from Faulkner County,
Arkansas, is not so far removed from the coastal plain, and may
have reached its site of collection via the Arkansas River also.
These three specimens from non-coastal plain localities are all
typical of the western subspecies, D. r. miaria.

Of the three subspecies of Deirochelys reticularia herein recog-
nized, D. r. miaria differs strikingly from the two eastern forms,
while both D. r. reticularia and D. r. chrysea are rather more closely
related. Differentiation on and along the Florida peninsula has
been demonstrated in many groups of animals, and likewise in many
genera and species of amphibians and reptiles. If degree of differ-
ence between subspecies is regarded as a criterion of length of time
required for differentiation and subspecies formation, it is apparent
that D. r. miaria has been isolated from D. r. reticularia longer than
has D. r. chrysea. It is possible that such isolation occurred during
glaciation, and that the Deirochelys population was at that time
split into two sections, one in Florida and southeastern North
America, and one in southern Texas and northern Mexico. During
this period of isolation, the characteristics of D. r. miaria were
intensified or developed, and with the return of more clement con-
ditions to the north, D. r. miaria invaded more northern and eastern
available areas. In the east, during the same period, differentiation
occurred on the varyingly extensive Florida peninsula, resulting in
the formation of D. r. chrysea. It seems preferable to regard both
D. r. chrysea and D. r. miaria as derivatives of D. r. reticularia.
The occurrence of a plastral blotch in some specimens of D. r.
reticularia may be interpreted as ancestral; on one hand, this plastral
marking has been amplified into the plastral pattern of D. r. miaria,

SCHWARTZ: CHICKEN TURTLE 499

and on the other has been completely lost in D. r. chrysea. The
latter subspecies also shows a significant reduction of the bridge
pattern and marginal markings in the bridge region, while such is
not the case in D. r. miaria. The shape of the shell of the three
subspecies is distinctive; the cuneiform shell of D. r. chrysea and the
flattened shell of D. r. miaria could have been developed most easily
from that of D. r. reticularia or its precursor.

The absence of Deirochelys from most of Mississippi and the
extension farther northward on the west side of the Mississippi
River are puzzling. It is possible that further collecting in Missis-
sippi will reveal the presence of this turtle in suitable localities in
that state and possibly also in westernmost Tennessee, although
it has not been taken in Reelfoot Lake, which may conceivably
be regarded as suitable habitat. The westernmost limit of the
distribution of the chicken turtle is the Balcones Escarpment,
and it should be sought in the area between the Escarpment and
its known range. Deirochelys is unknown from Mexico, and if it
occurs as far south as the Rio Grande that river may act as an
effective barrier to its further southern distribution. However, one
other southeastern pond-dwelling form (Siren intermedia) has been
taken in Tamaulipas, and the chicken turtle may occur at least
in that state.

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500

TABLE 3. COLORATION OF SHELL IN LIFE OF TEN SPECIMENS OF

Deirochelys reticularia
Specimens listed in the order noted in text

Carapace .

D. r. chrysea

. Chaetura Black (3)
black

Reticulum

. Yellow Ocher
Honey Yellow

Edge of carapace.

Plastron

. Ochraceous-Orange, blending to
Ochraceous-Buff posteriorly

Deep Chrome, blending to Light
Orange- Yellow posteriorly

. Antimony Yellow
Baryta Yellow centrally and
Deep Chrome peripherally

Bridge

. Orange with seams Empire Yellow
Light Orange Yellow

D. r. reticularia

Medal Bronze

black

black

black

black

black

Olive

Blackish Brown (3)

Citrine

Ochraceous-Tawny
Buckthorn Brown

(obscured)
Buffy Citrine
Ochraceous-Tawny
Buffy Olive
Honey Yellow

Pale Orange- Yellow
Yellow Ocher
Buff Yellow

(absent)
Old Gold
Yellow Ocher
Deep Colonial Buff
Reed Yellow

Pale Orange- Yellow,
grading to Orange-
Buff on sides of
gulars and humerals

Pale Orange-Yellow,
grading to Orange-
Buff on edges of gu-
lars and humerals
and lower sides of
anterior marginals

Maize Yellow

Buff-Yellow

Buff-Yellow

Maize Yellow

Cream-Buff

Naples Yellow

Orange-Buff, seams
Apricot Yellow

Maize Yellow, seams
Orange-Buff

Baryta Yellow

Baryta Yellow

Baryta Yellow

Capucine Orange

Cream-Buff

Naples Yellow

501

TABLE 4. COLORATION OF SOFT PARTS IN LIFE OF TEN SPECIMENS OF

Deirochelys reticularia
Specimens listed in the order noted in text

D. r. chrysea

Upper beak Wax Yellow

(not taken)

Dorsal head and
neck markings .

.Wax Yellow, grading posteriorly to

Old Gold

Picric Yellow, grading posteriorly
to Sulphine Yellow

Ventral neck

markings Martius Yellow

Martius Yellow and Pale Lemon
Yellow

Tail, hind limb,

and rump lines .... Martius Yellow
Picric Yellow

Forelimb stripe.

. Picric Yellow
Pale Lemon Yellow

D. r. reticularia

Martius Yellow
Picric Yellow
Martius Yellow
Martius Yellow
Martius Yellow
Picric Yellow
Sulphine Yellow
Citron Yellow

Pale Lumiere Green

Clear Dull Green- Yel-
low, grading to Olive
Ocher

Rainette Green

Pale Olivine

Glass Green

Pale Lemon Yellow

Warbler Green, grad-
ing to Sulphine
Yellow

Chalcedony Yellow,
grading to Old Gold

Light Greenish Yellow
Pale Greenish Yellow
Picric Yellow
Picric Yellow
Pale Greenish Yellow
Picric Yellow
Empire Yellow
Citron Yellow and
Massicot Yellow

Martius Yellow
Pale Greenish Yellow
Barium Yellow
Martius Yellow
Martius Yellow
Martius Yellow
Baryta Yellow
Barium Yellow

Pinard Yellow
Pale Greenish Yellow
Martius Yellow
Picric Yellow
Picric Yellow
Picric Yellow
Pinard Yellow
(not taken)

502

REFERENCES

CARR, ARCHIE F.

1952. Handbook of turtles. Comstock Publ. Assoc., Ithaca, N.Y., pp. xv,
1-542.

HARPER, FRANCIS

1940. Some works of Bartram, Daudin, Latreille, and Sonnini, and their
bearing upon North American herpetological nomenclature. Amer. Mid.
Nat., 23 (3), pp. 692-723.

LATREILLE, P. A.

1801. in SONNINI, C. S., and LATREILLE, P. A. Histoire naturelle des reptiles,
avec figures dessin^es d'apres nature, Paris, 1 : xx, 1-280.

RIDGWAY, ROBERT

1912. Color standards and color nomenclature. Washington, D.C., pp. iii,
1-43, 53 pis.

SCHMIDT, KARL P.

1953. A check list of North American amphibians and reptiles. Amer. Soc.
Ichth. and Herp., pp. viii, 1-280.

WEBB, ROBERT G.

1950. Range extension of the chicken turtle in Oklahoma. Herpet., 6 (5),
pp. 137-138.

WILLIAMS, ERNEST E.

1950. Variation and selection in the cervical central articulations of living
turtles. Bull. Amer. Mus. Nat. Hist., 94 (9), pp. 509-561.

WRIGHT, A. H., and FUNKHOUSER, W. D.

1915. A biological reconnaissance of the Okefinokee Swamp in Georgia. The
reptiles. Proc. Acad. Nat. Sci. Philadelphia, 67: 108-192, 14 text figs., 3 pis.

503

UNIVERSITY OF ILLINOIS-URBANA